‘\
ey
WN A canaie
Hen iade tate
(eid
wat
RA HN i aie
i re i
f
ars
ote as
Ty byt
Conway
‘Nt
i ey cay
ioe sh
Meruit x Ke) ati eh ait Dba!
CAM ely WIV Aavadn nen tolnatg
iin ta! Ne yi
es RAN Ay)
i
Carenniny,
WOVEN Ny
Wenn a!
Way LW ny.
wns WA
‘i saints
ies i
ca Mee Cat cette
Sethi
\
ve
etre all WAS
Varvdtay
We SA ade
SENN a
OM ath JValy.
¢ ene
: trent
Sas tata
mann SATURN
at aldaRet nto
anid u ik = ah N4Ciay WW sft at 14 mi
iow Te cots ea Ms Ah at
navi ehavia ha
ae
t a
r
aaa
Vicistahyne
ANS Lana
eh
av anenne dan ik
any An yan
Hitt
‘
Win say
wiley iARAY WW Ast Ay
stat hal Manny
yin
PM ay
SCL PR Rn ULY
PODS MET
HAS AD VEW Ny
ake Ni
WT utes Ny salty
NURM ANA us ae
AN LUN
WH Neate
hye
oN wenn
Serna er a ie Re
Dea MORO Anh eat
WM Wale he
Paranh
Ni meal
WOE CRIS Wawa
Seanad caine wa WeRby eyanntes
yay
Mitady
" be Pes Wee
Sa, Stemi eat en Seal ua
SEW WEG CEASA WOE ec Mh Vial aN MANN SiN iain a BENIN
wren ity " t. Sor Riri ats ch
nV WN Vy ays Deaton vn "
EES ! inane ta aaah nA Ue pumnaaans Ruts none
DME a BY iL a ay 43 Ney ete y NICSE HEM AN fri vst Es
SON oe Sty fiat ” ASM NY ¥s TAL wy at isthe
m vias ROR any 4 “y
CNW TE,
wy tela iy tas
YEO ra REA
MoV ities
ny
avatv bene
vn ys Daa “A
vy kan
Waban tis
saieiathangs my
REN ty hor TO dk tH S42 a te yi
vo vy tN statadh taka cea . ede sanunt Nace
Pee POMERAT et COM ere erer wey Ne
VD Acorn tiene i) alata vi alec
meta iMaN ey A TENA. an Wet)
Peg Ha thot Ss Mobo
Breer
LMR RIEL on
Wie yen
SPUN
AMSA Mts AQ Ny ein
IAN TY MSNA day IML Qe
dowate as ‘iil.
adits shag
DATES Serer
Sh MALIN
HOON DST
ITs DAWA
ees Aronian ir AAD RS
NWN Rates Aum gue R
BENG NAM MONE CHC MD
Wen SN WS HAR ETN QD
BWC
VoL Me, Ht WUT USAC HA ty
Tha CRC
CoS CRC Can Ct
AU en ie Be
VR Aas vial vey yey
Worn "
MIN INAES Vly VIS
AMONG Ra
witatbats
nrcucers ‘
hee Vey PR Ua eens
PIA AG a get We NYT 819
Vi tnd wy MIU Ad seen ita ewe H
on
Rinmennnee ft
DANI A
he SMT INL
Vena &
OSS Pn es DUNT Dry
RDC PON TOE nN
\ SAU PCT
SAMA Wy
Sabato
Cey
CTO ary
WIS Vea de an
n Nt SVN Le
at WATS
Vig
WHEE APR ON al alte
eek aay Wal mRSN
EY F wy
van
Pome)
NOPrIn wy
SSD SEU OTT)
He Sarin ata
Seti
AWA TI he
Aw
SRE AVON EN Renee.
COROT OT
Seren)
Vee pv
Howey
tee rae
IRR ace
MASA Aum,
athed od! OANPL AALS SITS
Vobev ML a eay
Va
seat
weafh chy hatin
AAS Syd
shes Ark
SS erat
VNR MTA LAU aaretes th
Awan
Wa Soden,
SARs
SONA
Ae
ys
gaye if *
' ‘ av Fak Sari)
ou va CoM ir aear rt
, , Pu lgs
‘ : yifenipn
. ' WP Sy meg SAP
‘ F 1pm ay
an Vy tom bap iiasy an
‘ . i Conor nny
é , ‘i rsh
ese ; en indies ath ey f
i A wi MT it heyy
be . ’ i aie nv 1 it a
. , ‘ , f
; 7 i Hit aii Laisa)
4 He
a Porte) tt. )
i ; 4 Hee elt airy ly
ue sz ” nig ‘ HA oy ay
ae wer roa . Sea tytn yw nytt Ae
r ‘ Pee chk pay oe
: ‘ fay att ER
. , Use
5 . tp ape
' coe i is i SERA es Fab ay eu peu td the Mies! a
; pee ' ae na ne ma Heaters tity
re tig His au
ie he : f i
ae ee ron se peikl “ay ic ceyH H
‘ ae a Se nung ae boned (a pi
. = ; wopeeny Di f fn Pi! Petre
‘ ‘ a pie Wie allen
' j Wah 9 ey en RAY
‘ ' ths Oh yale en
+ i 1 vv Woe ft Reo peek y Sey
; " iS (tea) ob 4 ite Hot hia al “f 2
, Aa as CAM OF ca tis rr Heiahs
A : ‘ ; ‘ SEW Miye eh ay athe ve wedi g Pan atite:
Dias he 7 eae jada Ry ftp sehanels
t 13 ri wi Ly vena Fee
a 1 i ifegngey H z CHU ne Hi A
‘ ‘ yh Hoge Oa. at Noyes na Seat vat iD sdb Pitt ta!
Y . , Vat unui Atay os i
‘ me it ‘ ue se Pan dawn anion y
soy fan ‘ torn
be 4 ' fb yWity H ays
1 : Mi AR ribs ls hetvannn by oy ei)
rf ‘ ior o f ety gigas) VINA in
oan cha cue? VA OE ME Ay te
: a sy.) Hi 40)
, Altice
Tai sine do pe : PRRCR
Lintoag t H A
VAs ican sas 7 ey
ve i ay i} at wy
vit Suites é Mand,
" “4 hos
aS
ae
¥
q
mau
y
ry : ’
A vy
if
4
aD
» 7 o ¥
, : I...)
Ty £ wa)
ion ay 4)
n
tel Yon" I
c “u
a ' 4s o
ral
DY eS ee
‘TT Lat vy
ty
4 /
. +
i ¥ i } '
' uy y
hf ai
eee |
er
i
'
4 ae
m
}
ie z f
7
, if
fi
i
ee
;
= iy
wd
a?
i ‘ ;
4 ! ran i
ih \ (,
en ul iV i
1 i |
i we ee a re ee
} { i i
hy Fj i A i
LM } , i f 4s } T) | -
any ‘alt ’ AM a oe 1 ie
ap sit! in ii ve hn wh
Fi TR OY Ak TR 1 ' } tt
i ( Whe) La | i
ay, inh) ihe RU ket ie MME Wels mar me
ny ed AON HAT FU en i A Yea 8 Ye, ii i
SNE tay Wy 4 Wey ir iy Carel | YN: de } Ait Wi A i
\ As, Y) ‘ ee! i Ly ae iL ety hie Wf NA Me a ae at Loe
; f i ne 4 vy Wao !
ae Rea Av Pas Pe ee me at We ey ial ea
ures nah ’ i i enh id f y f von ant an cee ry
te ayy ay ey i wy ‘ } ial ee ak Mate (|e yi ’ 1A ry j yh ih
; te Bh ve
i Van
a
AK
I Al
iss i"
q cas
na
——
it es
!
Ne
yi
iy)
ae
wi)
Se eet eM
Ns a | hiv i) ae) ;
le a f mo
la , :
' i : i he ,
Pe
pe , a Nie y ion
ori iM Lin it
Fae |
VOR; 99 JANUARY 1997 NO. 1
lo |
29K (ISSN 0013-8797)
Pai eh
PROCEEDINGS
of the
ENTOMOLOGICAL SOCIETY
of WASHINGTON
PUBLISHED
CONTENTS
BATRA, SUZANNE W. T.—Bionomics of Lasioglossum (Evylaeus) matianense (Blithgen)
(Hymenoptera: Halictidae), the predominant pollinating bee in orchards at high altitude in
ther GreatiHimalayajohGarhwall (UCP = india ace serrate oir tora Aare srattiegstet eee cles tee oteieten eee terse 162
COSCARON, MARIA DEL C. and JUAN J. MORRONE—Cladistics and biogeography of the
assassin bug genus Melanolestes Stal (Heteroptera: Reduviidae) ...................00..eeee eee Sys)
EVANS, HOWARD E. and STUART M. FULLERTON—Report on a collection of Bethylidae
@aymenoptera)iiroml central Hloriday 4 nrst oh wecloicrn vale ole ier erent satay enter 174
FITZGERALD, SCOTT J. and BORIS C. KONDRATIEFF—A new species of Pseudonomo-
neura Bequaert (Diptera. Viydidae) from Mexico a o.n acceso soueiece = erect ei en ee 171
GAGNE, RAYMOND J., CORNELIA OTT, and SUSANNE S. RENNER—A new species of
gall midge (Diptera: Cecidomyiidae) from Ecuador associated with flowers of Clavija (Theo-
(od a) EIST Yoho) | Rote aR ane A eRe Re RSME AG ATS GONE ceiind sot sao gem pHooUpoSad an nace orate oAdeHe 9: ‘i 110
GOEDEN, RICHARD D. and JEFFREY A. TEERINK—Notes on life histories and descriptions
of adults and immature stages of Procecidochares kristineae and P. lisae new species (Dip-
tera: Tephritidae) on Ambrosia spp. in southern California ..................0.ee ee eeeee eee ee ees 67
GOEDEN, RICHARD D. and JEFFREY A. TEERINK—Life history and description of imma-
ture stages of Procecidochares anthracina (Doane) (Diptera: Tephritidae) on Solidago cal-
iornrcauNuttall sin, Souther) Califormiaw y. 1. cerca. : seem soon eee eee ea etree te aie 180
KROMBEIN, KARL V. and BETH B. NORDEN—Nesting behavior of Krombeinictus nordenae
Leclercq, a sphecid wasp with vegetarian larvae (Hymenoptera: Sphecidae: Crabroni-
ip feet) Meta 0S a ol Sen as Aan RE Serene Oe Ati ear aaa Me tac AamA AK Gh tas nh acrid dan ion cide Sanam Adee 42
LOYA, LANE J. and J. E. MCPHERSON—Life history and laboratory rearing of Oedancala
dorsalis (Say) (Heteroptera: Lygaeidae), with descriptions of immature stages ............... 89
MACKAY, WILLIAM P.—A revision of the Neotropical ants of the genus Camponotus, sub-
eenus Myrmostenius (Hymenoptera: HOLMICIGAC)!Lovinsa as same nes islelee te need gelonsteeet = ace ress 194
MATILE, LOIC—Fenderomyia Shaw, a valid North American taxon in Macrocerinae (Diptera:
Mivcetophilordea mero attdae) i -.cietms cts aareetatsrs cle slas nlettatetclofeteinke eyetclelety eee tertiaae aii -efone aieios se) siete es 50
(Continued on back cover)
THE
ENTOMOLOGICAL SOCIETY
OF WASHINGTON
ORGANIZED MARCH 12, 1884
OFFICERS FOR 1997
M. ALMA SOLIS, President MICHAEL G. POGUE, Treasurer
WARREN E. STEINER, JR., President-Elect Davip G. FurtTH, Program Chair
HAROLD HARLAN, Recording Secretary Davip ADAMSKI, Membership Chair
HOLuis B. WILLIAMS, Corresponding Secretary RALPH P. ECKERLIN, Past President
Gary L. MILLER, Custodian
DAVID R. SmiTH, Editor
Publications Committee
THOMAS J. HENRY WAYNE N. MATHIS
Gary L. MILLER, Book Review Editor
Honorary President
CurRTIS W. SABROSKY
Honorary Members
LoulIsE M. RUSSELL ALAN STONE KARL V. KROMBEIN
All correspondence concerning Society business should be mailed to the appropriate officer at the following
address: Entomological Society of Washington, % Department of Entomology, MRC-168, Smithsonian Insti-
tution, Washington, D.C. 20560.
MEETINGS.—Regular meetings of the Society are held in the Natural History Building, Smithsonian Institu-
tion, on the first Thursday of each month from October to June, inclusive, at 8 P.M. Minutes of meetings are
published regularly in the Proceedings.
MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology.
Annual dues for members are $25.00 (U.S. currency).
PROCEEDINGS.—The Proceedings of the Entomological Society of Washington (ISSN 0013-8797) are pub-
lished quarterly beginning in January by The Entomological Society of Washington. POSTMASTER: Send
address changes to the Entomological Society of Washington, % Department of Entomology, MRC-168, Smith-
sonian Institution, Washington, D.C. 20560. Members in good standing receive the Proceedings of the Ento-
mological Society of Washington. Nonmember U.S. subscriptions are $60.00 per year and foreign subscriptions
are $70.00 per year, payable (U.S. currency) in advance. Foreign delivery cannot be guaranteed. All remittances
should be made payable to The Entomological Society of Washington.
The Society does not exchange its publications for those of other societies.
PLEASE SEE PP. 615-616 OF THE JULY 1996 ISSUE FOR INFORMATION 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, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash-
ington, D.C. 20560.
Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology,
Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560.
Books for Review: Gary L. Miller, Systematic Entomology Laboratory, ARS, USDA, Building 046, BARC-
West, Beltsville, MD 20705.
Managing Editor and Known Bondholders or other Security Holders: none.
This issue was mailed 25 February 1997
Second Class Postage Paid at Washington, D.C. and additional mailing office.
PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA
This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 1—27
ERETMOCERUS HALDEMAN (HYMENOPTERA: APHELINIDAE) IN THE
UNITED STATES, WITH DESCRIPTIONS OF NEW SPECIES ATTACKING
BEMISIA (TABACI COMPLEX) (HOMOPTERA: ALEYRODIDAE)
MIKE ROSE AND GREGORY ZOLNEROWICH
Department of Entomology, Texas A&M University, College Station, TX 77843, U.S.A.
Abstract.—A key to species of Eretmocerus that occur in the continental United States
is provided. Eretmocerus eremicus, n. sp., E. joeballi, n. sp., E. staufferi, n. sp., and E.
tejanus, n. sp., all reared from Bemisia (tabaci complex) collected in the United States,
are described. Redescriptions of Eretmocerus californicus Howard, E. corni Haldeman,
E. haldemani Howard, and E. portoricensis Dozier are provided. Eretmocerus debachi
Rose and Rosen, FE. furuhashii Rose and Zolnerowich, and E. illinoisensis Dozier are
discussed. Lectotypes are designated for Eretmocerus californicus Howard, E. haldemani
Howard, and E. portoricensis Dozier.
Key Words:
Species of Eretmocerus Haldeman, 1850
(Hymenoptera: Chalcidoidea: Aphelinidae)
are attracting widespread interest because
of population explosions of Bemisia (tabaci
complex) (Homoptera: Aleyrodidae) around
the world (Rose et al. 1996). All known
species of Eretmocerus are primary para-
sites of whitefly, and species of Eretmoce-
rus have been purposefully utilized in ef-
fective biological control programs (Rose
and DeBach 1991-1992; Rose and Rosen
1991-92; Rose 1988). This paper, which is
part of a larger study of Eretmocerus spe-
cies of the world, is designed to clarify
characterizations of named species in the
U.S., and to describe new species that at-
tack Bemisia (tabaci complex) in the U.S.
Because of confusion regarding the use
of the names Bemisia tabaci (Gennadius)
A-strain, B. tabaci B-strain, and B. argen-
tifolii Bellows and Perring on specimen la-
bels and in the literature (Brown et al.
1995), we refer to hosts from this group as
Bemisia (tabaci complex). Species of Be-
misia (tabaci complex) have emerged as a
Eretmocerus, Aphelinidae, Bemisia, Aleyrodidae, biological control
major agricultural pest attacking a variety
of food, cash, and ornamental crops in the
United States and abroad. Bemisia (tabaci
complex) attacks over 500 plant species in
74 families (Mound and Halsey 1978;
Brown and Bird 1992) and new host plants
are continually being added (Gill 1992;
Costa et al. 1993). Crop damage occurs due
to feeding, honeydew production and resul-
tant sooty mold, and disease transmission.
Species in the Bemisia (tabaci complex)
transmit more than 15 viruses (Byrne et al.
1990) that cause more than 40 plant dis-
eases (Brown and Bird 1992). Reflecting
the current U.S. situation, Bemisia (tabaci
complex) is a major pest of crops in Mex-
ico, the Caribbean, Central and South
America, the Middle East, India, and parts
of Africa.
Biological control research to attain pop-
ulation regulation of Bemisia (tabaci com-
plex) in the U.S. is emphasizing evaluation
of naturally occurring species of parasitic
Hymenoptera and importation of exotic
populations and species of parasites. The
2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
majority of native and imported species are
found in Eretmocerus and Encarsia Foers-
ter (Hymenoptera: Chalcidoidea: Aphelini-
dae). Polaszek et al. (1992) provided a key
and discussion of Encarsia species that at-
tack Bemisia (tabaci complex). Schauff et
al. (1996) furnished a pictorial key and dis-
cussion of species of Encarsia that attack
whitefly in North America. Species of Ami-
tus Haldeman (Hymenoptera: Platygastri-
dae) also have been reared from Bemisia
(tabaci complex) (Viggiani and Evans
1992):
Surveys to obtain samples of Bemisia
(tabaci complex) and its parasites from var-
ious host plants in the U.S. have been un-
dertaken primarily in Arizona, California,
Florida, and Texas. These states have suf-
fered the greatest damage to field crops and
subsequent losses to Bemisia (tabaci com-
plex), so biological control research efforts
have been emphasized in these areas. It is
vitally important that researchers in these
and other areas are able to identify the bi-
ological entities they are discovering, im-
porting, colonizing and evaluating.
There are currently 8 described species
of Eretmocerus from the New World, and
30 species are known from the Old World.
Given that there are nearly 1200 described
species of whitefly, and that species of Er-
etmocerus are known from all continents
where whitefly occur, there are undoubtedly
many more undescribed species of Eret-
mocerus. Because most species of Eretmo-
cerus have been reared from agricultural
pests, little is known of the actual range of
these species. Likewise, little is known
about species found outside of agricultural
settings.
The literature encompassing the taxono-
my of Eretmocerus is disparate in its com-
prehensiveness, and researchers have relied
on historical taxonomy to provide names
for numerous species of Eretmocerus that
are currently being studied and transferred
nationally and internationally. These histor-
ical names often find their way into new
literature and are then generally adopted,
which may obscure the actual species being
studied. For example, two of the species de-
scribed in this paper were called E. califor-
nicus or E. sp. nr. californicus for many
years.
Previous studies (Rose and Rosen 199]—
1992, Rose and Zolnerowich 1994, Rose et
al. 1996, Hunter et al. 1996) have empha-
sized developing criteria for the character-
ization of species based on morphology,
pigment patterns of males, reciprocal mat-
ing trials, and electrophoretic analysis.
These studies all indicate that minor mor-
phological differences in the size, shape, or
chaetotaxy of the habitus, antennae, and
forewings, and pigment patterns of males,
are significant species characters.
Males of different species of Eretmoce-
rus are very similar unless they have fea-
tures such as distinct pigment patterns. The
poor condition of male specimens available
for Eretmocerus haldemani and E. illinois-
ensis preclude their separation in the key to
species. Males are unknown for Eretmoce-
rus joeballi and E. portoricensis.
METHODS
High quality microslide mounts are nec-
essary to correctly examine and _ identify
species of Eretmocerus. Because clearing
and mounting specimens in balsam removes
all or most of their color, most specimens
used in this study were mounted in Hoyer’s
medium as described by Rosen and DeBach
(1979), and the cover slips were sealed with
two coats of red GLPT, a nonconducting
insulating varnish used in electronics (GC
Electronics, Rockford, Illinois). Other spec-
imens were critical point-dried and mount-
ed on cards prior to mounting in balsam.
Most primary type material was mounted in
balsam.
Position of the body and antenna on mi-
croslides is critical for proper identification.
The antenna should be examined from the
lateral aspect. Viewing the antenna from
any other angle obscures the true length and
shape of the segments. In many instances,
the antennae were removed from the head
VOLUME 99, NUMBER 1
fll
—_
ro
fl
1 2
3
Figs. 1-3. 1, 2 antenna. 2, Forewing. 3, Venation of forewing. c = length of club, fl = length of first
funicular segment, fll = length of second funicular segment, 1 = length of forewing, m = length of marginal
vein, p = length of pedicel, r = length of radicle, s = length of scape, sm = length of submarginal vein, st =
length of length of stigmal vein, w = greatest width of disc, wl = width I.
and mounted separately on the same slide.
Wings should be positioned away from the
body to reveal their shape and allow accu-
rate measurements to be made.
Length of the body was measured from
critical point-dried specimens using an eye-
piece micrometer when possible. Other
measurements of specimens mounted in
balsam or Hoyer’s were taken using an eye-
piece micrometer or a digitizing tablet. The
maximum length of each antennal article in
lateral view was measured (Fig. 1). The an-
tennal ratios given in descriptions are the
ratios of the lengths of the radicle : scape :
pedicel : club. Unless expressed as a range,
the length : width ratios represent average
values.
Length and width of the forewing were
measured as shown in Fig. 2. The maxi-
mum length (1) and width of the wing
across the disc (w) were measured. Width I
(wl) is the distance between the distal end
of the frenal fold and the anterior margin
of the forewing above the distal end of the
stigmal vein. Points of measurement for the
lengths of the submarginal, marginal, and
stigmal veins are shown in Fig. 3. The ra-
tios of the lengths of the longest anterior
and posterior alary fringes (Fig. 5) to the
width of the wing are given, as is the num-
ber of tubercles (Fig. 11), which are present
on the ventral side of the wing.
In many instances, the distal boundary of
the marginal vein and the advent of the stig-
mal vein were not clearly delimited. In
those cases, the end of the marginal vein
and the beginning of the stigmal vein were
taken at the base of the distal large seta on
the marginal vein (Fig. 3, st).
Descriptions of color for critical point-
dried and specimens mounted in Hoyer’s
may differ because of differences in speci-
£
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Species of Eretmocerus in the U.S. and original collection data.
Species Author Year Original Host Insect
corni types lost Haldeman 1850 Tetraleurodes corni (Haldeman)
[= Aleurodes corni Haldeman]
corni neotype Dozier 1932 Trialeurodes packardi (Morrill)
[= Trialeurodes morrilli (Britton)]
californicus Howard 1895 unknown ‘‘Aleyrodes’”’
haldemani Howard 1908 Aleuroplatus coronata (Quaintance)
[= Aleyrodes coronata Quaintance]
illinoisensis Dozier 1932 unknown
portoricensis Dozier 1932 Aleurothrixus floccosus (Maskell)
debachi Rose & Rosen 1991-92 Parabemisia myricae (Kuwana)
furuhashii Rose & 1994 Parabemisia myricae (Kuwana)
Zolnerowich
eremicus Rose & 1997 Bemisia (tabaci complex),
Zolnerowich possibly Trialeurodes
Joeballi Rose & 1997 Bemisia (tabaci complex)
Zolnerowich
staufferi Rose & 1997 Bemisia (tabaci complex),
Zolnerowich Trialeurodes abutiloneus (Haldeman)
tejanus Rose & 1997 Bemisia (tabaci complex)
Zolnerowich
' Host plant probably Quercus (R. Gill, personal communication).
men preparation. Specimens mounted in
Hoyer’s do not show colors as well as crit-
ical point-dried material, but do show fus-
cous pigment patterns well. Descriptions of
color or pigment are given for both critical
point-dried and Hoyer’s-mounted speci-
mens where appropriate.
For new and described species, label data
for primary types is presented exactly as
recorded on the specimen labels, with the
data for each individual label enclosed by
quotation marks and each line of the label
separated by a slash. Label data for material
other than primary types are standardized
and multiple records from the same locality
may be combined. Table | provides sum-
mary information such as original locality,
host, and host plant for the species of Er-
etmocerus discussed here.
The following acronyms represent insti-
tutions or individuals who loaned material
for study, are repositories for type material,
or are otherwise mentioned in the text:
BMNH, The Natural History Museum,
London, United Kingdom; CDFA, Califor-
nia Department of Food and Agriculture,
Sacramento; HU, The Hebrew University,
Rehovat, Israel; INHS, Illinois Natural His-
tory Survey, Champaign; MJR, personal
collection of M. J. Rose; NSM, National
Science Museum, Tokyo, Japan; TAMU,
Texas A&M University, College Station;
UCR, University of California, Riverside;
USNM, National Museum of Natural His-
tory, Smithsonian Institution, Washington,
Dic
KEY TO SPECIES OF ERETMOCERUS IN THE
UNITED STATES
1 Female, antenna with 2 funicular seg-
moe (Yt EMTS, I) ae ac occa dcaon DZ
_ Male, antenna without funicular seg-
THENUS SS. hoe oe sence eee esse ate ces 1
2(1) Mesosoma brown to brown orange, con-
trasting strongly with head and metasoma
(Fase 25) i. x teartupceettays E. staufferi, n. sp.
- Body yellow, mesosoma not contrasting
with head and metasoma ............ 3
3(2) Mesoscutum with 4 setae (Fig. 15) ..... 4
VOLUME 99, NUMBER I
iablewle
Continued.
Nn
Original Host Plant
Original Locality, Date
Cornaceae: Cornus sericea, dogwood
Balsaminaceae: /mpatiens biflora, jewel weed
Fagaceae: Quercus agrifolia, California live oak
unknown!
unknown
Burseraceae: Bursera simaruba
Zygophyllaceae: Guaiacum officinale
Rutaceae: Citrus spp.
Moraceae: Morus sp., mulberry
Malvaceae: Gossypium hirsutum, cotton
Moraceae: Morus sp.
Solanaceae: Solanum
Solanaceae: Lycopersicon, tomato
Cruciferae: Brassica oleracea, cabbage
Pennsylvania, 1847
Pennsylvania, White Clay
Creek, 1929
California, Los Angeles, 1887
California, Berkeley, 1908
Illinois, Elizabethtown, 1932
Puerto Rico: Bayamon &
Central Aguirre, 1925
California, Orange County,
1982
Japan: Honshu, 1979
Arizona, Phoenix, 1991
California, Bakersfield, 1993
Texas, College Station, 1993
Texas, Mission, 1994
5(3)
6(5)
7(6)
Mesoscutum with 6 setae
Parapsis with 2 setae; club 4.1—4.5 times
as long as its greatest width (Fig. 8)
BAe ARI Ree Se es, Le La E. debachi
Parapsis with 3 setae (Fig. 15); club 4.3-
5.6 times as long as its greatest width (Fig.
13) E. furuhashii
Club 3.2-3.8 times as long as wide, with
dorsal surface convex and contrasting with
straight ventral surface (Fig. 20)
Betts Pitas ton ses LEIS, SePEEITSE E. portoricensis
Club 4.5—8.3 times as long as wide, and
with dorsal and ventral surfaces more or
less parallel (e.g., Figs. 4, 26)
Dorsal surface of club apically tapered so
that apex forms a broad point (Fig. 4); rad-
icle 0.6 times or more as long as scape
E. californicus
Apex of club truncate; radicle 0.35—0.50
times as long as scape (e.g., Figs. 16,
26)
Pedicel short, 2—2.3 times as long as wide;
club 4.5—5.1 times as long as wide (Fig.
18); ovipositor 1.5 times as long as club
and 1.3 times as long as midtibia
E. joeballi, n. sp.
Pedicel longer, 2.5—4 times as long as
wide; club 5.9-8.3 times as long as wide
(e.g., Figs. 6, 16); ovipositor shorter than
or equal to length of club or midtibia ... 8
9(8)
10(9)
11(1)
12(11)
Scape at least 6.5 times as long as wide
(Fig. 16); midbasitarsus 8.5 times as long
as wide E. haldemani
Scape 3.5—5.5 times as long as wide (e.g.,
Fig. 10); midbasitarsus 5.5—8.3 times as
long as wide
Pedicel 2.4—3.1 times as long as wide, and
0.25—0.3 times as long as club (Fig. 6);
reared from Trialeurodes ........ E. corni
Pedicel 3—4 times as long as wide, and
0.29—0.39 times as long as club (Fig. 10);
reared from Bemisia (tabaci complex) and
possibly Trialeurodes
Gastral tergite II usually with | pair, but
occasionally 2 pairs, of lateral setae (Fig.
12); gastral sternites anterior to base of
Ovipositor usually with a group of 4—5 se-
tae, occasionally with 3—6 setae
3 sat gd SRE sciyiyl ne cme umes aaa as E. eremicus, 0. sp.
Gastral tergite II usually with 2 pairs, but
occasionally 1 pair, of lateral setae (Fig.
28); gastral sternites anterior to base of
Ovipositor usually with a group of 7-8 se-
tae, occasionally with 5—6 setae
BO, Ora Ere Ps cna roe ae E. tejanus, n. sp.
Mesoscutum with 4 setae (Fig. 15) .... 12
Mesoscutum with 6 setae
Parapsis with 2 setae; mesoscutum with
dark fuscous “‘T’’ shape; scutellum com-
pletelyatuscouss(Biga29) pa -eaee- E. debachi
6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
= Parapsis with 3 setae (Fig. 15); mesoscu-
tum with light fuscous *““T’’ shape; scutel-
lum medially fuscous, lateral areas unpig-
isacsynieerol (Vente SN) 55555505000 E. furuhashii
13(11) Entire mesosoma brown to brown orange,
contrasting strongly with head and meta-
GYopmnkey (IFW, BY) ooo ou aod E. staufferi, n. sp.
- Mesosoma not brown to brown orange, al-
though fuscous pigment patterns on pro-
notum, mesoscutum, or scutellum may be
present or absent (e.g., Figs. 30,33) ... 14
14(13) Mesoscutum with longitudinal fuscous
bandsi(Hic=33) ie oe. o E. tejanus, n. sp.
- Mesoscutum without longitudinal fuscous
bands cc BWR ne Goac. Gas seems IS)
15(14) Mesoscutum fuscous along anterior
margin (Fig. 30), or with a light fuscous
“‘T” shape (as in Fig. 31), or evenly fus-
COWS! i 2e5 as ee eee een oar 16
- Mesoscutum not fuscous............ 197;
16(15) Hind tibia fuscous ...... E. eremicus, n. sp.
- Hind tibia not fiscous) 2-25-24. - E. corni
17(15) Scape 2.5—2.8 times as long as pedicel .. .
MN Ds ate ee ar eRe Lc, hepa ls E. californicus
- Scape 3.2—4 times as long as pedicel .. .
ENC ele ice ours E. haldemani, E. illinoisensis
Eretmocerus californicus Howard
(Figs. 4—5)
Eretmocerus californicus Howard, 1895: 16
[USNM, examined].
Type material. Lectotype here designat-
ed, 2 mounted in balsam on a slide la-
belled, ‘“‘Name Eretmocerus/californicus/
Howard/REMOUNT/Det HOWARD 19
1895/Coll D. W. Coquillett/No. USNM 2699/
Com" LECTOTYPE? "Ci" “toc. Tos
Angeles/California/Date VI-9 19 1887/Host
Aleyrodes/Det 19/On Quercus/agrifolia’’.
There are an additional five 2 and 16 d
paralectotypes mounted on 15 slides
[USNM].
Diagnosis. Females of E. californicus can
be distinguished by the length of the radi-
cle, which is 0.6 or more than the length
of the scape, the extremely reduced first fu-
nicular segment, and the tapered shape of
the club (Fig. 4). Eretmocerus californicus
is most similar to E. eremicus and E. teja-
nus, which differ in having the radicle only
0.4—0.5 the length of the scape, the first
funicular segment is not as reduced, and the
club is not tapered (Figs. 10, 26).
Males of E. californicus lack pigment
patterns on the mesoscutum, and the scape
is 2.5-2.8 X as long as the pedicel. Eret-
mocerus eremicus and E. tejanus differ by
having distinct fuscous pigment patterns on
the mesoscutum (Figs. 30, 33).
Female.—Length and body color could
not be accurately determined from available
slide-mounted specimens. Wings hyaline.
Face and occiput with transverse substri-
gulate sculpture, interscrobal area vertically
substrigulate. Antenna (Fig. 4) with radicle
5.4X as long as wide; scape 3.9X as long
as wide, 1.5—1.6X length of radicle, 1.6X
length of pedicel, 0.5 length of club; ped-
icel 2.6X as long as wide, slightly shorter
than radicle, 0.6X length of scape, 0.3
length of club. Funicle I triangular, 1.2 as
wide as long and much smaller than funicle
II; funicle II 1.8 as wide as long. Club
with apex apically tapered, 5.2—6.6X as
long as wide, 2X length of scape, 3.2
length of pedicel. Antennal ratio 1:1.5:1.0:
Sole
Mesoscutum trapezoidal with 6 setae and
with reticulate sculpture in the anterior %,
remainder with elongate reticulate sculpture
laterally and substrigulate sculpture medi-
ally. Parapsis with 2 setae and faint reticu-
late to substrigulate sculpture; axilla with 1
seta and sculpture similar to parapsis. Scu-
tellum with 4 setae and 2 placoid sensilla
lateral to and close to posterior setae, lat-
erally with elongate reticulate sculpture, re-
mainder with elongate to substrigulate
sculpture. Propodeum smooth. Endophrag-
ma extending to posterior half of gastral ter-
gite II.
Forewing (Fig. 5) 2.7X as long as wide
at width I, 2.4 as long as maximum width
of disc. Longest anterior alary fringe 0.2
width of disc, longest posterior alary fringe
0.4X< width of disc. Single seta at base of
wing present or absent; distal portion of
costal cell with 2—5 setae. Marginal vein
with 3 long setae, 10—12 setae usually be-
tween marginal vein and linea calva. Linea
VOLUME 99, NUMBER 1
4
/
af
SS
oo "
~ N= mets ae tS
ERR a aay ee
2 —~ ~ a ~
SOE BRS SN es & ~\
mae ane ~ Sim) ~ =
in ee SS
$a 4 Sa ~ \ ~~ > TES; =
a ae ea YN NY ~ Va SS
es ee aa eng eit CN We \
= ie e
ed ae a ~\s\ ply Vw Sens
See SE Sats \, EN
E ee SAIN ee Teal aa! j
— aN Ne Sade Sey J a?
me oe , Ss
Re ees Bae Say Nan ANG OD Coe
~*~ eet e\ VN S e
ad Sis Z Nees
Ry aoa SS by SS
Proc | Sy ay
pr 7 = eye aS
eee —— x
ie SEEN
Set iy, ee
db ee
GA? mn _ ry, es
CF rN a AEE Gi
Si AE EB re ee
— mer =
- Fe eerie —
; Cee ee
sz - = ——— ae
DSI IE (OTN mn 1 a Vas eee, Gp am Nw ee
oz vr 3 me
7 cd (if 4 EDN RE =
; { a a
- dy Tne Kacey pe ee He
/ Ly hd “¢ Fi i Te aol
= =
eS ghia ihbee8t eS oe NS ee
Mr 55 x Ly rr Sj eas
‘ Sear aS we
ca Pfr sare =
ee
72 re
or eo 4
: — rm :
OE as Sal a
_ [eg ae rm ap %&
A 4 LV wn oo
Wo i —- pe te ean
a, ee y, a a
~~~ 2S mea le ee \ yy S a
= —
3 i, - apa UF ?
r=) art Hi, oo erie PN
Sa, Bente hwy, SOS GN erst a er
MA v fr i¢ = — _ ~
2 “7 Umeete Eyes ERS eS
a2 °
= Fs $e ~ CN eta
LU TMG oS ei oe
- t —
oy
Sc SS
2
Figs. 4-9. 4-5, Eretmocerus californicus. 4, 2 antenna. 5, 2 forewing. 6—7, E. corni. 6, 2 antenna. 7, 2
forewing. 8-9, E. debachi. 8, 2 antenna. 9, 2 forewing. af = anterior alary fringe, pf = posterior alary fringe.
calva closed posteriorly by setae, with 12—
15 tubercles on ventral surface of wing near
posterior end of linea calva; a group of 31—
43 setae including those forming distal edge
of linea calva point toward anterior margin
of wing, remaining 171—175 setae in disc
point to distal apex of wing. Ratio of sub-
marginal:marginal:stigmal veins 2.8:1.1:1.
Hind wing 6.9 as long as wide, with 7—
9 setae in center.
Gastral tergite I covered with strong re-
ticulate sculpture; lateral margins of gastral
tergites imbricate with dense stippling; gas-
tral tergites I-VI with paired setae as fol-
lows: 1, 1, 2, 2-3, 2, 1. Syntergum with 4
setae.
8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Ovipositor slightly exserted and equal to
length of club, 3X length of pedicel, 1.9
length of scape, and 1.1 length of midtib-
ia.
Male.—Body color could not be accu-
rately determined from available slide-
mounted specimens. Mid- and hind tarsi I—
III fuscous. Marginal and stigmal veins fus-
cous, base of forewing and costal cell
slightly fuscous.
Host.—Reared from an unknown species
of Aleyrodes.
Discussion.—Howard’s original descrip-
tion and figures of E. californicus were
based on a series of female and male spec-
imens reared from an undetermined Aley-
rodes collected by D. W. Coquillett in Los
Angeles, California, in 1887. Although
Howard (1895) deposited type material in
the USNM, and Dozier examined the ma-
terial in 1932, Compere (1936) was unable
to examine E. californicus and placed it in
his key based on the original description.
The type material was lost from about 1932
until 1992.
In 1992, G. Evans found the missing
slides of E. californicus at the USNM. The
specimens on one slide (five female, eight
male) were crumpled in dark orange balsam
and not suitable for study. All the speci-
mens on this slide have been individually
remounted in balsam. G. Evans also re-
mounted broken specimens, including one
female, from two other slides labelled as
USNM Type No. 2699. The label data for
these two new slides, which list “‘Astero-
diaspidis”’ as the host, do not agree with the
label data from the other type slides. We
consider this to be an error.
Our figures and redescription are derived
from the six remounted females and eight
remounted males, and seven male speci-
mens from a single original Howard slide.
There are now a total of 16 slides bearing
E. californicus specimens labelled with the
original collection data and identified as
USNM Type No. 2699.
It is interesting to note that FE. californi-
cus was reared from an unknown species of
Aleyrodes on oak. Aleuroplatus, Tetraleu-
rodes, and Trialeurodes all occur on Quer-
cus in southern California. M. Rose and J.
B. Woolley (TAMU) were unable to obtain
Eretmocerus from numerous collections of
whitefly, particularly Tetraleurodes stanfor-
di (Bemis), from oaks in Los Angeles
County over the past 15 years.
Lack of accurate characterization of E.
californicus has resulted in misapplication
of this name, particularly to species of Er-
etmocerus reared from Bemisia (tabaci
complex) in the U.S. We have not examined
any specimens of E. californicus reared
from Bemisia (tabaci complex).
Eretmocerus corni Haldeman
(Figs. 6-7)
1850:
Eretmocerus corni Haldeman, 110
[USNM, examined].
Type Material—Neotype ¢ mounted in
balsam on a slide labelled. ““Eretmocerus
corni Hald./Reared from white-/fly, Aster-
ochiton/sp. on Impatients/biflora./White
Clay Creek/Pa. Sept. 16, 1929/H.L Dozier”
‘“‘Eretmocerus/corni Hald./Designated as/this
long-lost/species./2 + ¢ Neotype/U.S.N.M.”
Diagnosis.—Females of Eretmocerus
corni can be distinguished by the combi-
nation of the elongate club that is 5.9-7X
as long as wide and the relatively short ped-
icel, which is only 2.4—3.1X as long as
wide, and 0.25—0.29X, rarely 0.30X, as
long as the club (Fig. 6). Eretmocerus corni
is similar to E. eremicus and E. tejanus,
which differ in having the pedicel 3—4X as
Jong as wide and 0.28—0.39% as long as the
club (Figs. 10, 26). Eretmocerus haldemani
also has an elongate club, but it is 7.5-8.3X
as long as wide (Fig. 16).
Males of Eretmocerus corni have the me-
soscutum and scutellum fuscous, and the
hind tibia is not fuscous. They are most
similar to males of E. eremicus and E. te-
janus. Males of E. eremicus differ in having
the hind tibia fuscous, and males of E. te-
janus can be distinguished by the longitu-
VOLUME 99, NUMBER 1
dinal fuscous bands on the mesoscutum
(Fig. 33).
Female.—Length of specimens in Hoy-
er’s 0.47—0.55 mm. Specimens in Hoyer’s
with body and antennae pale yellow. Head
amber; eyes amber with inner red pigment,
ocelli pale. Wings hyaline.
Face and occiput with transverse substri-
gulate sculpture, interscrobal area vertically
substrigulate. Antenna (Fig. 6) with radicle
4x as long as wide; scape 4.9 as long as
wide and 2.1X length of radicle, 2.2X
length of pedicel, 0.4 length of club; ped-
icel 2.4—3.1X as long as wide, 0.4 length
of scape, 0.25—0.29X, and rarely 0.30,
length of club. Funicle I triangular, 1.25
as wide as long; funicle II subquadrate.
Club with apex truncate, 5.9—7X as long as
wide, 2.5 length of scape, 4.9 length of
pedicel. Antennal ratio 1:2.1:1.6:4.7.
Mesoscutum trapezoidal with 6 setae and
reticulate sculpture anteriorly, remainder
with elongate reticulate sculpture. Parapsis
with 2 setae and reticulate to substrigulate
sculpture; axilla with | seta and reticulate
to substrigulate sculpture. Scutellum with 4
setae and 2 placoid sensilla lateral to and
slightly closer to posterior setae than to an-
terior setae, with fine substrigulate sculpture
medially, remainder with reticulate sculp-
ture. Propodeum medially reticulate, faintly
reticulate on lateral margins. Endophragma
extending nearly to posterior margin of gas-
tral tergite IV.
Forewing (Fig. 7) 2.9X as long as wide
at width I, 2.7 as long as maximum width
of disc. Longest anterior alary fringe 0.16
width of disc; longest posterior alary fringe
0.3X width of disc. Single seta at base of
wing present or absent; distal portion of
costal cell usually with 3, occasionally 2—
5, setae. Marginal vein with 3 long setae;
10-11 setae, occasionally 10—15, between
marginal vein and linea calva. Linea calva
closed posteriorly by setae, with 12—16 tu-
bercles on ventral surface of wing near pos-
terior end of linea calva; a group of 32—45
setae including those forming distal edge of
linea calva point toward anterior margin of
wing, remaining 183-238 setae in disc
point to distal apex of wing. Ratio of sub
marginal:marginal:stigmal veins 3:1.3:1.
Hind wing 7.5X as long as wide, with 7—
15 setae in center.
Gastral tergite I with faint transverse sub-
strigulate sculpture medially and reticulate
sculpture on anterior half of lateral margins;
lateral margins of tergites II-VI imbricate
with stippling; tergites I-VI usually with
paired setae as follows: 1, 1, 2, 2, 2, 1, but
occasionally 1, 1—3, 2—3, 2—3, 2, 1. Synter-
gum with 4 setae.
Ovipositor slightly exserted, 0.8 length
of club, 3.4 length of pedicel, 1.8 length
of scape, equal in length to midtibia.
Male.—Specimens mounted in balsam
with head orange. Apex of scape, pedicel,
and multiporous plate sensilla fuscous. Pro-
notum fuscous. Mesoscutum fuscous,
darkest along anterior margin and becoming
paler posteriorly; tegula slightly fuscous.
Scutellum fuscous, darkest along anterior
and posterior margins. Metanotum and pro-
podeum slightly fuscous. Aedeagus slightly
fuscous. Fore- and hind tarsi slightly fus-
cous; distal portion of midtibia and midtarsi
I-III fuscous, midtarsus IV less fuscous.
Submarginal vein, costal cell, and edges of
marginal and stigmal veins fuscous; base of
forewing and venation of hind wing slightly
fuscous.
Host.—Reared from Trialeurodes pack-
ardi (Morrill), the strawberry whitefly.
Discussion.—Haldeman’s (1850) original
description and figure of the type species,
E. corni, were based on “‘Two mutilated
specimens”’ reared from Tetraleurodes cor-
ni (Haldeman) [= Aleurodes corni Halde-
man]. Unfortunately, Haldeman did not
designate a depository for his specimens.
Dozier (1932), who was unable to locate
the original material, provided a redescrip-
tion of the species and designated a neotype
female based on a series of specimens (15
2, 3 3) reared from Trialeurodes packardi
(Morrill) [as 7. morrilli (Britton)] and an
additional field-collected female.
The slide labelled as the neotype lists
10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
‘‘Asterochiton”’ as the host, but Dozier only
mentions Trialeurodes morrilli as a host in
the redescription. Mound and Halsey
(1978) do not list any species of Asterochi-
ton in the U.S. Dozier’s original identifica-
tion of the host is probably an error. Al-
though Dozier stated the slide bearing the
neotype female also contained one addition-
al female and one male specimen, the slide
with the red USNM type label only has a
single female and single male. Despite
these discrepancies, we accept this slide
with the red USNM type label as the one
with the neotype.
Eretmocerus corni did not successfully
reproduce on Bemisia (tabaci complex) in
limited trials at TAMU. It may prove to be
important in the biological control of spe-
cies of Trialeurodes.
Other material examined.—USA: Mary-
land: Blackwater Refuge, reared from 7.
packardi on wild strawberry, 19.x, 24,
26.x.1942, H. L. Dozier (1 2, 2 6, USNM);
Cambridge, 24, 26, 30.vili., 9.1x.1943, Ex.
Trialeurodes on Euphorbia hirsuta, H. L.
Dozier (1 2, 6 6, USNM); Church Creek,
Ex. Trialeurodes on Lizard’s Tail, 2,
Axe 94). sit OE tad SINM): = Church
Greek:) Ex, Lizard?s: Vail}, 101942) (io
USNM); New York: Ithaca, Cornell Plan-
tations, 29.ix.1992, Trialeurodes packardi
on Impatients pallida, T92047, G. W. Fer-
rentino (8 2, 4 ¢, MJR); Ithaca, S. of Bee-
be Lake, 10.viii.1988, ex. Trialeurodes
packardi, M. Hunter (1 6, MJR).
Eretmocerus debachi Rose and Rosen
(Figs. 8-9, 29)
Eretmocerus debachi Rose and Rosen,
1991-1992: 200 [USNM, examined].
Type material.—Holotype 2 mounted in
balsam on a slide labelled, “‘Name Eret-
mocerus &/debachi/Holotype sp.nov./Bal-
sam/Det Rose 1982/Coll M. Rose/No.
B2.76/Corr. USNM/Rose&Rosen 91-92”
‘‘Loc Rancho Mission/Viejo, Orange Co.,
Cal./Date VII.15 1982/Host Parabemisia/
myricae/Det Rose 1982/On Valencia or-
ange/and lemons’”’.
There are five 2 and one d paratypes
with the same label data as the holotype.
These are deposited with the BMNH, HU,
MJR, TAMU, and USNM.
Diagnosis.—Females of E. debachi can
be distinguished by the deflexed apex of the
club (Fig. 8), mesoscutum with 4 setae, and
each parapsis with 2 setae. It is most similar
to E. furuhashii, which differs in having a
longer club (Fig. 13) and 3 setae on each
parapsis (Fig. 15). Eretmocerus joeballi can
have the apex of the club slightly deflexed
(Fig. 18) but differs in having 6 setae on
the mesoscutum.
Males of E. debachi have each parapsis
with 2 setae, a fuscous ““T’’ shape on the
mesoscutum, and the scutellum is com-
pletely fuscous (Fig. 29). Males of E. fu-
ruhashii differ in having each parapsis with
3 setae, a very light fuscous ““T”’ shape on
the mesoscutum, and the lateral areas of the
scutellum are unpigmented (Fig. 31).
Host.—Reared from Parabemisia myri-
cae (Kuwana), the bayberry whitefly.
Discussion.—The description of E. de-
bachi was based on series of specimens
reared from Parabemisia myricae collected
on citrus in southern California. The tax-
onomy, biology and history of E. debachi
in California is well documented (Rose and
DeBach 1991-1992, Rose and Rosen 1991—
92, Rose and Zolnerowich 1994). This spe-
cies has been introduced and established in
Israel and Turkey, where it is responsible
for control of P. myricae. Introductions are
planned for Spain and France (J. C. Onil-
lon, personal communication) and Morocco
(D. Rosen, personal communication).
Eretmocerus eremicus Rose and
Zolnerowich, new species
(Figs. 10—12, 30)
Diagnosis.—Females of E. eremicus can
be distinguished by the club that is 6.5—
7.3X as long as wide (Fig. 10), pedicel that
is 3—4X as long as wide and 0.3—0.39X as
VOLUME 99, NUMBER 1
Figs. 10-12.
long as the club, mesoscutum with 6 setae,
and gastral tergite II usually with 1 pair, but
occasionally 2 pairs, of lateral setae (Fig.
12):
Females of E. eremicus are very similar
to E. corni and E. tejanus. Females of E.
corni have the pedicel 2.4—3.1 as long as
wide and 0.25—0.30 as long as the club,
and have not been reliably recorded from
Bemisia (tabaci complex). Eretmocerus te-
janus has the anterior alary fringe of the
forewing slightly shorter (Fig. 27), and gas-
tral tergite II usually has 2 pairs, but oc-
casionally 1 pair, of lateral setae (Fig. 28).
Males of E. eremicus have the meso-
scutum fuscous along the anterior margin
(Fig. 30), or with a light fuscous ““T”’ shape
(as in Fig. 31), or evenly fuscous, and the
hind tibia is fuscous. Males of E. corni do
not have the hind tibia fuscous, and males
of E. tejanus can be distinguished by the
SS SS SSN
SS Sey > ~~
Si “N EN oe ‘N
~
SSS Sore os
SSE SR SSSR VX
ERR RS RES TSS NR 4, —
Soa TR Were \ 8S
Se St eS es? <I
ee =a N \ Ny Doline Now ramen ®
pee ~ Wi bcicciene
— * .
11
Eretmocerus eremicus. 10, 2 antenna. 11, 2 forewing. 12, 2 gaster. t = tubercle.
longitudinal fuscous bands on the mesocu-
tum (Fig. 33).
Female.—Length of critical point-dried
specimens 0.45—0.55 mm. Body light yel-
low. Head light yellow or yellow white and
paler than body; eyes grey green, ocelli red.
Scape same color as face, pedicel and fla-
gellum slightly darker. Legs pale yellow
and paler than body, tarsal claws dark.
Wings hyaline.
Specimens mounted in Hoyer’s with
body and legs pale yellow amber, legs oc-
casionally slightly darker than body. Head
pale to light amber. Antenna pale amber.
Wing venation and frenal fold of forewing
amber.
Face and occiput with transverse substri-
gulate sculpture, interscrobal area vertically
substrigulate. Antenna (Fig. 10) with radi-
cle 4.2X as long as wide; scape 5.1X as
long as wide, 2.1 length of radicle, 1.6
1)
length of pedicel, 0.6 length of club; ped-
icel 3-4 as long as wide, 1.3 length of
radicle, 0.6 length of scape, 0.3—0.39x
length of club. Funicle I triangular, 1.1 as
wide as long; funicle II 1.1X as long as
wide. Club 6.5—7.3X as long as wide, 1.7 X
length of scape, 2.6 length of pedicel. An-
tennal ratio 121 -1.3:3°5:
Mesoscutum trapezoidal and with 6 se-
tae, occasionally with supernumary setae,
anterior 4%—% with reticulate sculpture, me-
dially with substrigulate sculpture, remain-
der with elongate reticulate sculpture. Par-
apsis with 2 setae and substrigulate sculp-
ture; axilla with | seta and substrigulate
sculpture. Scutellum with 4 setae and 2
placoid sensilla lateral to and closer to pos-
terior setae or equidistant, with substrigu-
late sculpture in median area, remainder
with elongate reticulate sculpture. Propo-
deum with substrigulate sculpture. Endo-
phragma extending to posterior half of gas-
tral tergite IV.
Forewing (Fig. 11) 2.9 as long as wide
at width I, 2.7 as long as maximum width
of disc. Longest anterior alary fringe 0.17
width of disc, longest posterior alary fringe
0.35 width of disc. Base of wing usually
with 1, occasionally O—2, setae, distal por-
tion of costal cell usually with 2-3 setae.
Marginal vein with 3 long setae, 6—12 setae
between marginal vein and linea calva. Li-
nea calva closed posteriorly by setae, with
11—14 tubercles on ventral surface of wing
near posterior end of linea calva; a group
of 24—35 setae including those forming dis-
tal edge of linea calva point toward anterior
margin of wing, remaining 120—202 setae
in disc point to distal apex of wing. Ratio
of submarginal:marginal:stigmal veins 3.5:
leseit
Hind wing 6.9X as long as wide and with
5—11 setae in center.
Gastral tergite I with strong substrigulate
sculpture anterolaterally, remainder substri-
gulate; lateral margins of tergites faintly im-
bricate with stippling; gastral tergites I-IV
(Fig. 12) usually with paired setae as fol-
2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
lows: 1, 1, 1,,1,.2,.1, but cam be:l,-1—2, 1—
2, 1-2, 2, 1. Syntergum with 4 setae.
Ovipositor slightly exserted, equal in
length to club, 2.6 length of pedicel, 1.7
length of scape, equal in length to midtibia.
Male.—Length of critical point-dried
specimens 0.43—0.51 mm. Face light yel-
low, vertex orange yellow, eyes grey green,
ocelli red. Scape same color as face or
slightly darker, pedicel and flagellum tes-
taceous. Pronotum dorsolaterally brown.
Mesoscutum orange yellow to orange
brown and fuscous along anterior margin
(Fig. 30), or with a light fuscous “‘T’’ shape
(as in Fig. 31), or evenly fuscous; parapsis
lighter. Scutellum colored as mesoscutum.
Dorsellum brown. Propodeum light brown
dorsally. Gaster yellow brown to orange
brown dorsally, remainder yellow. Coxae
pale whitish, proximal %—% of all femora
pale white, remainder tan to light brown, all
tibiae light brown, all tarsi brown.
Specimens mounted in Hoyer’s with head
dark amber. Radicle and scape fuscous;
pedicel darker, club slightly fuscous and
darker at the base, multiporous plate sen-
silla dark fuscous. Pronotum fuscous. Me-
soscutum fuscous along anterior margin
(Fig. 30), or with a light fuscous ““T”’ shape
(as in Fig. 31), or evenly fuscous; parapsis
and axilla occasionally slightly fuscous.
Scutellum usually fuscous, darkest along
anterior and posterior margins. Metanotum
medially fuscous. Propodeum fuscous,
darker along lateral margins. Gastral ter-
gites slightly fuscous. Aedeagus fuscous.
Foreleg beyond coxa fuscous, tibia darkest;
midleg beyond coxa fuscous, tarsi I-III
darkest; hind leg beyond coxa fuscous, tarsi
IV lighter than proximal tarsi. Base of fore-
wing and costal cell slightly fuscous; ve-
nation of fore- and hind wings fuscous.
Host.—Reared from Bemisia (tabaci
complex) and possibly Trialeurodes. Spec-
imens have been reared from mixed hosts
of Bemisia and Trialeurodes, with the latter
most likely 7. abutiloneus (Haldeman), the
bandedwinged whitefly, or 7. vaporariorum
(Westwood), the greenhouse whitefly.
VOLUME 99, NUMBER 1
Discussion.—This species has been re-
ferred to as E. californicus, E. sp. nr. cali-
fornicus, and E. haldemani. Although fe-
males of E. eremicus are extremely similar
to those of E. tejanus, differences in male
pigment patterns, in conjunction with recip-
rocal mating tests and electrophoretic anal-
ysis between populations from Arizona and
California, and Texas (Hunter et al. 1996),
indicate that the Arizona and California
populations form a species distinct from the
population in Texas.
Eretmocerus eremicus is the most com-
mon naturally occurring parasite of Bemisia
(tabaci complex) in the desert areas of Ar-
izona and California. This species possibly
parasitizes Trialeurodes abutiloneus (Hal-
deman), which can be sympatric with Be-
misia in those areas. Headrick et al. (1995,
1996) described the behavior of E. eremi-
cus.
Etymology.—From Latin
meaning “‘of the desert’’.
Holotype.— mounted in balsam on a
slide labelled, “‘Loc Phoenix/Arizona/Date
III 1991/Host Bemisia/tabaci/Det GB 1991/
On Gossypium/hirsutum”’ “‘Name Eretmo-
cerus/@/spm 1/7? USNM/Det Rose 1991/Coll
G. Butler/No. GB-3/Corr. Rose/Noyes 91”’.
Deposited in USNM.
Paratypes.—USA: same data as holotype
(3 2 and 3 6 mounted in balsam, 19 2 and
7 3 mounted in Hoyer’s); Texas, Hidalgo
Co., Mission Biological Control Labora-
tory, F, lab culture, M94001, original ma-
terial: USA: California, Imperial Co.,
Brawley, K. Hoelmer, 6.1.1994, ex: Bemisia
tabaci, on: okra (4 2 and 2 ¢ mounted in
balsam, 7 2 and 4 ¢ card-mounted). Para-
types deposited with BMNH, MJR, TAMU,
UCR, and USNM.
Other specimens examined.—USA: Ar-
izona: Maricopa Co., Phoenix, greenhouse,
23.iv.1992, Bemisia tabaci on collards, O.
Minkenberg (2 2, 5 3); Pima Co., Tucson,
21.x.1991, Bemisia tabaci on cotton, O.
Minkenberg (10 2, 10 d); Tucson, green-
house, 11.vi.1993, Bemisia sp. on pointset-
tia, O. Minkenberg (6 @, 13 d); Califor-
eremicus,
13
nia: Imperial Co., 16.xi.1992, Bemisia sp.,
i. Perrine. (2.5228 3,6 UCR); Bard,
7.v1.1993, Bemisia sp. on Cucurbita sp., K.
Hoecimere (io 2, 13 6); Brawley;
29.x1.1982, Bemisia tabaci, cotton, Kramer
& Forrester (11 2,7 3); Brawley, 8.v.1982,
Bemisia & Trialeurodes on \antana, G. But-
ler (16 2, 12 3); Brawley, 21.viii.1982, Be-
misia tabaci on cotton, Rose (10 2, 10 3);
Brawley, 29.xi.1983, Bemisia tabaci, cot-
ton, Kramer & Forrester (5 2, 2 6); Braw-
ley, 23.x.1984, Bemisia tabaci, bindweed,
D. Meyerdirk (19 @, 3 6); Brawley,
USDA, 13.1x.1993, kenaf, Rose & Pickett
(5 2,4 3); Brawley, 8.v.1995, Bemisia ar-
gentifolii on melon in refuge, M. Rose (10
2); Brawley, Legion Rd./Hwy. 86,
11.14.1995, W. Roltsch, Bemisia argentifolii
on kenaf (19 2, 1 6); Brawley, Legion Rd./
Hwy. 86, 20.iv.1995, Bemisia argentifolii
on eggplant, W. Roltsch (18 @); Brawley,
Legion Rd/Hwys 86, I8-v-1995, WwW:
Roltsch, Bemisia argentifolii on cantaloupe
(3 2); Brawley, Legion Rd./Hwy. 86,
22.v.1995, Bemisia argentifolii on sunflow-
er, W. Roltsch (14 2, 1 6); Brawley, Le-
gion Rd./Hwy. 86, 13.vi.1995, W. Roltsch,
Bemisia argentifolii on collard (14 2, 1 3);
Brawley, Legion Rd./Hwy. 86, 19.vii.1995,
W. Roltsch, Bemisia argentifolii on cotton
(9 2); Brawley, Legion Rd./Hwy. 86,
21.vili.1995, W. Roltsch, Bemisia argenti-
folii on cotton (24 2); El Centro, Desert
Trails RV Park, 1.11.1993, Bemisia on cot-
ton, M. Rose (3 2, 1 6); El Centro, Desert
Trails RV Park, 1.11.1993, Bemisia on Hi-
biscus, Rose & Pickett (12 2, 11 6); Holt-
ville, 13.ix.1993, Bemisia on sunflower, M.
Rose (32 2, 105 ¢); nr. Holtville, CDFA-
Bornt’s, 13.ix.1993, Bemisia on sunflower,
Rose, Pickett, & Roltsch (33 2, 18 3);
Kern Co., Bakersfield, 8.x.1992, Trialeu-
rodes abutilonea? on cotton, J. Ball (2 @);
Bakersfield, 1.1x.1993, Bemisia tabaci on
eggplant, J. Ball (i) 9):5 Bakersfield,
1.x.1993, Bemisia tabaci on lantana, J. Ball
(1 2, 1 o); Bakersfield, 29.vi.1994, Bemi-
sia on lantana, J. C. Ball (3 ° ); Bakersfield,
16.viii.1994, Bemisia on lantana, J. Ball (6
14 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
B sKe™ ~
‘ a9
15
Figs. 13-15.
mesoscutum, p = parapsis, s = scutellum.
2,8 3); Bakersfield, 17.viii.1994, Bemisia
on okra, J. Ball (6 2, 2 ¢d); Bakersfield,
7.1x.1994, Bemisia on fruitless mulberry, J.
Ball (2 2, 1 3); Bakersfield, 8.1x.1994, Be-
misia on lantana, Joe Ball (1 d). All spec-
imens from MJR unless noted otherwise.
Eretmocerus furuhashii Rose and
Zolnerowich
(Figs. 13-15, 31)
Eretmocerus furuhashii Rose and Zolne-
rowich, 1994: 286 [NSM, examined]
Type material—Holotype 2 mounted in
balsam on a slide labelled, “‘Name Eret-
mocerus/furuhashii/sp.nov./Det 19/Coll M.
Rose/No./Corr.”’ “‘Loc Japan:/Shizuoka,
Kochi/Date VIII.1979/Host Parabemisia/
myricae (Kuwana)/Det 19/On Morus.”
There are an additional 198 paratypes
from a number of localities in California
and Japan mounted in balsam and Hoyer’s
14
Eretmocerus furuhashii. 13, 2 antenna. 14, 2? forewing. 15, 2 mesonotum. a = axilla, m =
on a number of slides. These are deposited
with BMNH, HU, MJR, NSM, TAMU, and
USNM.
Diagnosis.—Females of E. furuhashii
can be distinguished by the deflexed apex
of the club (Fig. 13), mesoscutum with 4
setae, and each parapsis with 3 setae (Fig.
15). It is most similar to E. debachi, which
differs in having a shorter club (Fig. 8) and
only 2 setae on each parapsis. Eretmocerus
joeballi can have the apex of the club
slightly deflexed (Fig. 18) but differs in
having 6 setae on the mesoscutum.
Males of E. furuhashii have each. parap-
sis with 3 setae, a light fuscous ““T’’ shape
on the mesoscutum, and the lateral areas of
the scutellum are unpigmented (Fig. 31).
Males of E. debachi differ in having each
parapsis with 2 setae, a dark fuscous ““T”’
shape on the mesoscutum, and the scutel-
lum is completely fuscous (Fig. 29).
VOLUME 99, NUMBER 1
// 18
Eretmocerus haldemani. 16, 2 antenna. 17, 2 forewing. 18-19, E. joeballi. 18, 2 antenna. 19,
Figs. 16-19.
2 forewing.
Host.—Reared from Parabemisia myri-
cae, the bayberry whitefly.
Discussion.—The description of EF. fu-
ruhashii was based on specimens reared
from Parabemisia myricae collected on
mulberry in Japan and citrus in southern
California (Rose and Zolnerowich 1994).
This species was introduced to California in
1979 and was established in citrus groves
in Orange, Riverside, and Ventura counties
prior to the rapid distribution of E. debachi
(Rose and DeBach 1991-1992). No recent
evidence of E. furuhashii has been found in
southern California, but displacement may
not be complete in all areas of citriculture.
Eretmocerus haldemani Howard
(Figs. 16—17)
Eretmocerus haldemani Howard, 1908: 65
[USNM, examined].
Type material.—Lectotype here desig-
nated, 2 mounted in balsam on a slide la-
19
belled, “Morrill No. 509/? holotype/Type/
No. 11708/U.S.N.M.”’ “‘Eretmocerus n.sp./
2 /haldemani/How.”
An additional slide bears two d speci-
mens, labelled “Morrill No. 509/Type/NO.
11708/U.S.N.M.”’ ‘‘Eretmocerus/nsp. d/
haldemani How.”’ These are designated as
paralectotypes.
Diagnosis.—Females of E. haldemani
can be distinguished by the extremely elon-
gate club that is 7.5—-8.3 as long as wide
(Fig. 16), the elongate scape that is 6.9X as
long as wide, and the elongate midbasitar-
sus that is 8.5 as long as wide. It is most
similar to E. corni, which has an elongate
club that is only 5.9—-7X as long as wide,
and the scape is only 4.9X as long as wide
(Fig. 6).
Males of Eretmocerus haldemani are
very similar to those of E. illinoisensis.
Males of these species cannot be diagnosed
or separated based on the poor condition of
specimens available for study.
16 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Female.—Length, body color, sculpture,
and setation could not be accurately deter-
mined from the available slide-mounted
specimen. Wings hyaline.
Antenna (Fig. 16) with radicle 4.2 as
long as wide; scape 6.9X as long as wide,
2.5 length of radicle, 1.8 length of ped-
icel, 0.6X length of club; pedicel 3.6 as
long as wide, 1.4X< length of radicle; 0.5 x
length of scape, 0.3 length of club. Fu-
nicle I triangular, 1.2 as long as wide; fu-
nicle II subquadrate. Club elongate, apex
truncate, 7.5—-8.3X as long as wide, 1.7X
length of scape, 3.2 length of pedicel. An-
tennal ratio 1:2.5:1.3:4.4.
Mesoscutum trapezoidal and with 6 se-
tae. Parapsis with 2 setae; axilla with | seta.
Scutellum with 4 setae and 2 placoid sen-
silla lateral to and closer to posterior setae
than anterior setae.
Forewing (Fig. 17) 2.7X as long as wide
at width I, 2.4 as long as maximum width
of disc. Longest anterior alary fringe 0.09 x
width of disc, longest posterior alary fringe
0.25 width of disc. Base of wing with 1—
2 setae, distal portion of costal cell with 4—
5 setae. Marginal vein with 3 long setae,
14-16 setae between marginal vein and li-
nea calva. Linea calva closed posteriorly by
setae, with 16 tubercles on ventral surface
of wing near posterior end of linea calva; a
group of 44—48 setae including those form-
ing distal edge of linea calva point toward
anterior margin of wing, remaining 221 se-
tae in disc point to distal apex of wing. Ra-
tio of submarginal:marginal:stigmal veins
2.4:1:1.
Hind wing 7X as long as wide, with 13—
14 setae in center.
Male.—Body color could not be accu-
rately determined from available slide-
mounted specimens. Foretarsi and mid- and
hind tarsi IV slightly fuscous; mid- and
hind tarsi I-III fuscous. Base of costal cell
and venation of forewing fuscous.
Host.—Reared from Aleuroplatus coro-
nata (Quaintance), the crown whitefly.
Discussion.—Howard (1908) described
E. haldemani from one female and one
male reared from Aleuroplatus coronata
(Quaintance) collected by E. M. Ehrhorn in
Berkeley, California. No collection dates
are given in the literature or on the type
slides. Although Howard stated that he de-
scribed this species from one female and
one male, the two slides bearing red USNM
type labels bear one female and two males,
respectively. Despite this discrepancy we
accept the slides with the USNM type la-
bels as being Howard’s type specimens. No
host plant was recorded, but A. coronata is
common on Quercus in northern California.
Numerous specimens of Eretmocerus
held in collections in the U.S. and abroad
have been erroneously identified as E. hal-
demani. M. Rose, K. Hoelmer (USDA/
APHIS), and C. Pickett and R. Gill (CDFA)
have made numerous unsuccessful attempts
over the past 15 years to rear specimens of
Eretmocerus from Aleuroplatus coronata
collected on oaks in Berkeley, California.
Eretmocerus illinoisensis Dozier
Eretmocerus illinoisensis Dozier, 1932: 114
[INHS, examined].
Type material—Holotype ¢d mounted in
balsam on a slide labelled *“‘Eretmocerus/
illinoisensis/d Dozier/Swept from veg-/eta-
tion in bed of/creek./Elizabethtown, II1./
Aug. 5-1932/H.L. Dozier’ ‘‘Eretmocerus/
illinoisensis/Dozier/SL.12934 I.N.H.S./
Type No.” ‘‘HOLOTYPE/Eretmocerus/il-
linoisensis/DOZIER’”’ [on underside of
slide].
There are two additional d paratype
slides labelled ‘‘Eretmocerus/illinoisensis/d
Dozier/Swept from veg-/etation in bed of/
creek./Mounted in Euparol/Elizabethtown,
Ill./Aug. 5-1932/H.L. Dozier’ ‘‘Eretmoce-
rus/illinoisensis/Dozier/SL. 12935/Paratype
3 I.N.H.S./No.” ““PARATYPE/Eretmoce-
rus/illinoisensis/ DOZIER” [on underside of
slide], and ‘‘Eretmocerus/illinoisensis/d
Dozier/Swept from veg-/etation in bed of-
/creek./Elizabethtown, IIl./Aug. 5-1932/
H.L. Dozier’? *“‘Eretmocerus/illinoisensis/
Dozier/Paratype d/No. U.S.N.M.”
VOLUME 99, NUMBER I
Discussion.—Dozier’s (1932) description
of E. illinoisensis was based on a series of
six male specimens collected by sweeping
weeds and grasses in a creek bed near Eliz-
abethtown, Illinois. We were able to locate
only the holotype and two paratypes. Fe-
males are unknown, and Compere (1936)
was unable to include this species in his
key. As there are no host records, future
association with female specimens is prob-
lematic. Eretmocerus illinoisensis is very
similar to males of E. haldemani and cannot
be diagnosed or separated based on the poor
condition of specimens available for study.
Eretmocerus joeballi Rose and
Zolnerowich, new species
(Figs. 18-19)
Diagnosis.—Females of FE. joeballi can
be distinguished by the slightly deflexed
apex of the club (Fig. 18), 6 setae on the
mesoscutum, and the ovipositor is 1.5 the
length of the club and 1.3 the length of
the midtibia. It is most similar to Eretmo-
cerus debachi and E. furuhashii, which dif-
fer by having only 4 setae on the meso-
scutum (Fig. 15).
Female.—Length of specimens mounted
in Hoyer’s 0.7—-0.74 mm. Specimens in
Hoyer’s with head, body, and legs pale yel-
low. Portions of body with pupal exuviae
attached are gold orange. Wings hyaline.
Antenna (Fig. 18) with radicle 3.8 as
long as wide; scape 4.5 as long as wide,
2.1X length of radicle, 1.6 length of ped-
icel, 0.6X length of club; pedicel 2.0—2.3 x
as long as wide, equal in length to radicle;
0.5 length of scape, 0.3 length of club.
Funicle I triangular, 1.2 as wide as long;
funicle II 1.4 as wide as long. Club with
apex slightly deflexed, 4.5—5.1 x as long as
wide, 1.6X length of scape, 3.2 length of
pedicel. Antennal ratio 1:2.1:1:3.2.
Mesoscutum trapezoidal and with 6 se-
tae, anterior % with strong reticulate sculp-
ture, medially with substrigulate sculpture,
laterally with elongate reticulate sculpture.
Parapsis with 2 setae and strong substrigu-
17
late sculpture; axilla with 1 seta and sub-
strigulate sculpture. Scutellum with 4 setae
and 2 placoid sensilla lateral to and closer
to posterior setae, with substrigulate sculp-
ture, laterally with elongate reticulate sculp-
ture. Propodeum with faint substrigulate
sculpture medially, substrigulate sculpture
stronger around lateral margins. Endo-
phragma extending to posterior margin of
gastral tergite II.
Forewing (Fig. 19) 2.7X as long as wide
at width I, 2.4 as long as maximum width
of disc. Longest anterior alary fringe 0.14
width of disc, longest posterior alary fringe
0.3X width of disc. Base of wing usually
with 2-3 setae, distal portion of costal cell
with 2 setae. Marginal vein with 3 long se-
tae, 7-13 setae between marginal vein and
linea calva. Linea calva closed posteriorly
by setae, with 11—14 tubercles on ventral
surface of wing near posterior end of linea
calva; a group of 22-27 setae including
those forming distal edge of linea calva
point toward anterior margin of wing, re-
maining 87—149 setae in disc point to distal
apex of wing. Ratio of submarginal:margin-
al:stigmal veins 2.8:1.2:1.
Hind wing 6.7 as long as wide and with
1—4 setae in the center.
Gastral tergite I with substrigulate sculp-
ture, lateral margins with broad imbricate
sculpture and with stippling; gastral tergites
I-VI with paired setae as follows: 1, 1, 1,
2, 2, 1. Syntergum with 4—6 setae.
Ovipositor slightly exserted and 1.5
length of club, 4.6 length of pedicel, 2.3 x
length of scape, 1.3 length of midtibia.
Male.—Unknown.
Host.—Reared from Bemisia (tabaci
complex).
Discussion.—This uncommon species
has been recovered from Bemisia (tabaci
complex) on Morus and Solanum in Kern
County, California. Eretmocerus eremicus
occurs in the same area.
Etymology.—Named in honor of Joe
Ball, who collected the described speci-
mens and has contributed to biological con-
18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
=
3 a) Ves So
= <A Z is Oe Ree ea ee
a L Lye as aS
— ny era / pip Ls Pa = 2 oN ae
= =
a Ge Lag oe ge sfc
RCaGY, = awe ey
rf
22
Eretmocerus portoricensis. 20, 2 antenna. 21, 2 antenna with artifactual ‘parrot beak” club.
Figs. 20-22.
22, 2 forewing.
trol by natural enemies in California for
more than 35 years.
Holotype.—¢ mounted in Hoyer’s on a
slide labelled, ““Name Eretmocerus x/joe-
balli Rose +/Zolnerowich 96/2 @ °/Det.
Rose+Zoln. 1995/Coll.. J.. €C. Ball/No.
93-XI 22/Corr.”’ “‘Loc Bakersfield, CA/K-
ern Co./Date Oct. 1 1993/Host Bemisia Ta-
baci/Det 19/On Mulberry?’ There are two
? mounted on this slide. The holotype is
the uppermost specimen and delimited by
red ink. The second specimen on the slide
is desigated as a paratype. Deposited in
USNM.
Paratypes.—USA: same data as holotype
(2 2 mounted in Hoyer’s); California:
Kern Co., Bakersfield, 1.1x.1993, Bemisia
tabaci on nightshade (1 2 mounted in Hoy-
er’s); Bakersfield, 7.1x.1994, Bemisia on
fruitless mulberry, J. Ball (1 2 mounted in
Hoyer’s). Paratypes deposited with MJR
and TAMU.
Eretmocerus portoricensis Dozier
(Figs. 20-22)
Eretmocerus portoricensis Dozier, 1932:
115 [USNM, examined].
Type material—Lectotype here desig-
nated, 2 mounted in Hoyer’s on a slide la-
belled, ‘‘Eretmocerus/portoricensis/Dozier/
Reared from Aleu-/rothrixus flocco-/sus on
*“Almacigo”’/Bayamon, P Rico/Jan. 21-1925/
H.L. Dozier’ ‘‘Eretmocerus/portoricensis/
Dozier/?/Type. No. 44825 U.S.N.M.”
There are 13 2 specimens mounted on the
slide. The lectotype is the specimen closest
to the bottom of the slide, and faces the top
edge of the slide. It is partially circled with
black ink. The remaining specimens on the
slide are designated as paralectotypes.
An additional slide labelled *‘Eretmoce-
rus/californicus How./det Gahan/Reared
from/Aleurothrixus/floccosus Mask./on Al-
macigo/Bayamon, P.R./Jan. 21-1925/H. L.
VOLUME 99, NUMBER 1
Dozier” “‘Eretmocerus/portoricensis/Dozier/
Para//Type No. 44825 U.S.N.M.”’ has 19 @
specimens mounted on it. These are also
designated as paralectotypes.
We are unable to locate the other four
slides that Dozier cited in his original de-
scription.
Diagnosis.—Females of E. portoricensis
can be distinguished by the shape of the
club, which is strongly convex dorsally and
straight ventrally (Fig. 20). The shape of
this club is unlike that of any other de-
scribed species in the U.S.
Female.—Length of specimens mounted
in Hoyer’s 0.75—0.89 mm. Type series
mounted in balsam with body yellow, legs
slightly paler. Specimens in Hoyer’s with
body pale yellow. Wings hyaline.
Face and occiput with transverse substri-
gulate sculpture, lower face with substri-
gulate sculpture curving downward, inter-
scrobal area vertically substrigulate. Anten-
na (Fig. 20) with radicle 3.8X as long as
wide; scape 3.5 as long as wide, 2.1
length of radicle and pedicel, 0.7 length
of club; pedicel 2.2 as long as wide, equal
in length to radicle, 0.5 length of scape,
0.3 length of club. Funicle I triangular, as
wide as long; funicle II 1.6 as wide as
long. Club with apex curved, 3.2—3.8X as
long as wide, 1.5X length of scape, 3.1
length of pedicel. Antennal ratio 1:2.1:1:
Bul.
Mesoscutum trapezoidal and with 6 se-
tae, anterior 4 with reticulate sculpture, lat-
erally with faint elongate reticulate sculp-
ture, remainder with substrigulate sculpture.
Parapsis with 2 setae and faint reticulate to
substrigulate sculpture along lateral mar-
gins; axilla with | seta and sculpture similar
to parapsis. Scutellum with 4 setae and 2
placoid sensilla lateral to and closer to pos-
terior setae, with substrigulate sculpture
medially and reticulate sculpture laterally.
Propodeum with transverse striations me-
dially and reticulate sculpture laterally. En-
dophragma broad, extending from posterior
% of gastral tergite I to anterior margin of
gastral tergite II.
Forewing (Fig. 22) 2.7X as long as wide
at width I, 2.3X as long as maximum width
of disc. Longest anterior alary fringe 0.1
width of disc, longest posterior alary fringe
0.2 width of disc. Base of wing usually
with 2 setae, occasionally 1—3, often with 1
seta enlarged and reaching submarginal
vein, distal portion of costal cell usually
with 2—3, occasionally 1—4, setae. Marginal
vein with 3 long setae, 10—13 setae between
marginal vein and linea calva. Linea calva
closed posteriorly by setae, with 10—16 tu-
bercles on ventral surface of wing near pos-
terior end of linea calva; a group of 18—30
setae including those forming distal edge of
linea calva point toward anterior margin of
wing, remaining 139-191 setae in disc
point to distal apex of wing. Ratio of sub-
marginal:marginal:stigmal veins 3:1.3:1.
Hind-wing 6.9X as long as wide and
with 0-2 setae in center.
Gastral tergite I reticulate anteriorly and
anterolaterally; lateral margins of tergites
imbricate with rows of stippling; tergites
I-VI with paired setae as follows: 1, 1-2,
1-2, 1-2, 2—4, 2—4. Syntergum with 4 se-
tae.
Ovipositor slightly exserted, 1.2 length
of club, 3.6 length of pedicel, 1.7 length
of scape, equal in length to midtibia.
Male.—Unknown.
Host.—Reared from Aleurothrixus floc-
cosus (Maskell), the woolly whitefly.
Discussion.—Dozier’s description was
based on a series of 45 female specimens
on three slides reared from Aleurothrixus
floccosus (Maskell) on Bursera (= Ela-
phrium) simaruba (L.) taken at Bayamon,
Puerto Rico, and 18 female specimens on
three slides reared from A. floccosus on
Guaiacum (= Guajacum) officinale L. col-
lected at Central Aguire, Puerto Rico.
Gahan originally identified Dozier’s
specimens as E. californicus in 1925. Do-
zier (1932) later compared his specimens
from A. floccosus with the type material of
E. californicus at the USNM and concluded
the species were distinct. The antennal club
of E. portoricensis was described with “the
20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
apex shaped somewhat like a parrot’s
beak’’, but this distinctive shape is artifac-
tual. The apex of the club is a blunt curve,
but the shriveled contents of the club can
withdraw from the cuticle to create a
beaked appearance (Fig. 21). This artifact
has not been observed in other species.
Eretmocerus portoricensis has been
reared from A. floccosus collected on citrus
in Peru. This material probably resulted
from the importation of parasites of A. floc-
cosus from Florida in 1957 (Ebeling 1957).
Other specimens examined.—USA:
Florida: Broward Co., Fort Lauderdale,
ix.10.80, Aleurothrixus floccosus on citrus,
R. Dowell, No. R80-52 (13 2); Fort Lau-
derdale, 1982, Aleurothrixus floccosus, C.
R. Thompson (7 @); Hollywood, ii.2.80,
Aleurothrixus floccosus on citrus, Wm.
Gregory., No. R80-1-2, No. R80-1-3 (21
2); Hollywood, viii.31.81, Aleurothrixus
floccosus on citrus, Wm. Gregory, No.
R81-45 (6 2); Oakland Park, 1.3.80, Aleu-
rothrixus floccosus on citrus, B. Gregory,
No. R80-1-11 (3 2); Manatee Co., Anna
Maria Island, 20.11.1993, Aleurothrixus floc-
cosus (Mask.) on sea grape, Rose & Kring
(7 2); PERU: Lima: ix.1974, Aleurothrixus
floccosus on citrus, J. S. Torres (9 @). All
specimens from MJR.
Eretmocerus staufferi Rose and
Zolnerowich, new species
(Figs. 23-25, 32)
Diagnosis.—Females of E. staufferi can
be distinguished by the extremely elongate
club that is 8.2—9.1x as long as wide (Fig.
23) and the distinct brown or brown orange
mesosoma (Fig. 25). No other species has
such an elongate club or the mesosoma
brown orange.
Males also have the distinct brown or
brown orange mesosoma (Fig. 32) that is
unlike that of any other species of Eret-
mocerus.
Female.—Length of critical point-dried
specimens 0.53—0.63 mm. Critical point-
dried material with vertex orange red; face
and gena paler and more yellow. Antenna
colored as face. Mesosoma brown orange.
Gaster yellow orange. Legs varying from
testaceous to tan, tarsi slighter darker than
tibiae. Wings slightly fuscous. Specimens
mounted in Hoyer’s usually with mesoscu-
tum and scutellum pale medially (Fig. 25).
Face and occiput with transverse substri-
gulate sculpture, interscrobal area vertically
substrigulate. Antenna (Fig. 23) with radi-
cle 4.3X as long as wide; scape 5.7X as
long as wide, 2.5 length of radicle, 1.8
length of pedicel, 0.6 length of club; ped-
icel 3.3 as long as wide, 1.4 length of
radicle, 0.6 length of scape, 0.3 length
of club. Funicle I triangular, 1.2 as long
as wide; funicle II 0.9X as long as wide.
Club 8.2—9.1 X as long as wide, 1.8 length
of scape, 3.3X length of pedicel. Antennal
ratio 1:2.5:1.4:4.5.
Mesoscutum trapezoidal, usually with 6
setae but supernumerary setae common, an-
terior %4%—% with strong reticulate sculpture,
narrow median area with substrigulate sculp-
ture, remainder with elongate reticulate sculp-
ture. Parapsis with 2 setae and substrigulate
sculpture; axilla with | seta and substrigulate
sculpture. Scutellum with 4 setae and 2 plac-
oid sensilla lateral to and closer to posterior
setae, with strong substrigulate sculpture, me-
dially varying from reticulate to substrigulate,
posterolateral margins varying from reticulate
to substrigulate. Propodeum with substrigu-
late sculpture medially, laterally with strong
substrigulate sculpture, remainder smooth or
slightly substrigulate. Endophragma extend-
ing to posterior margin of gastral tergite I.
Forewing (Fig. 24) 3X as long as wide
at width I, 2.7 as long as maximum width
of disc. Longest anterior alary fringe 0.17
width of disc, longest posterior alary fringe
0.3 width of disc. Base of wing usually
with 3—4, occasionally 2—6, setae, distal
portion of costal cell usually with 2—3, oc-
casionally 1—4, setae. Marginal vein with 3
long setae, 8-12 setae between marginal
vein and linea calva. Linea calva closed
posteriorly by setae, with 10—17 tubercles
on ventral surface of wing near posterior
VOLUME 99, NUMBER 1
25
Figs. 23-25.
end of linea calva; a group of 33—39 setae
including those forming distal edge of linea
calva point toward anterior margin of wing,
remaining 126—223 setae in disc point to
distal apex of wing. Ratio of submarginal:
marginal:stigmal veins 2.9:1.2:1.
Hind wing 7X as long as wide and with
16—38 setae in the center.
Gastral tergite I with substrigulate sculp-
ture, lateral margins with broad imbricate
sculpture and with stippling; gastral tergites
I-VI with paired setae as follows: 2—6, 3—
7, 3-7, 4-8, 4-10, 3—4. Syntergum with 5—
10 setae.
Eretmocerus staufferi. 23, 2 antenna. 24, 2 forewing. 25, 2 habitus.
Ovipositor slightly exserted and 0.8
length of club, 2.6 length of pedicel, 1.4
length of scape, 0.9 length of midtibia.
Male.—Specimens mounted in Hoyer’s
with head amber; eyes bright red. Radicle
and scape slightly fuscous, scape darker in
distal %; pedicel dark fuscous; multiporous
plate sensilla fuscous. Pronotum fuscous.
Mesoscutum fuscous, darkest anteriorly
(Fig. 32). Parapsis, axilla, tegula, and scu-
tellum fuscous, scutellum darkest along
margins. Metanotum fuscous, darkest me-
dially. Propodeum fuscous, darkest along
lateral margins. Gaster slightly fuscous,
22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
with gastral tergite I darker medially. Ae-
deagus fuscous. Foreleg except coxa and
trochanter slightly fuscous; midtrochanter,
femur, and tibia slightly fuscous, tarsi
darker; hind leg slightly fuscous, femur and
proximal % of tibia darkest. Fore- and hind
wings slightly fuscous.
Host.—Reared from Bemisia (tabaci
complex) and Trialeurodes abutiloneus
(Haldeman), the bandedwinged whitefly.
Discussion.—There were only two male
specimens in the original collections of E.
staufferi. Males are not common in labo-
ratory cultures and are not needed for pro-
duction of females. This distinct thelyto-
kous species is being evaluated for its bio-
logical control potential in California, Tex-
as, and the Netherlands.
Etymology.—Named in honor of R. S.
Stauffer, who found this species attacking
Bemisia (tabaci complex) and T. abutilo-
neus at TAMU.
Holotype.—? mounted in balsam on a
slide labelled, ““USA: Texas/Hidalgo Co./
Mission/Biological/Control/Laboratory/F,
lab culture/M94002”’ ‘‘original/material:/
USA: Texas/Brazos Co./College Station/
10.xii.1993”’ “Rose & Stauffer/ex: Bemi-
sia/tabaci/on: tomato’’. Deposited in
USNM.
Paratypes.—USA: same data as holotype
(6 2 mounted in balsam, 15 2 card-mount-
ed); Texas: College Station, glasshouse,
TAMU, 29.xi.1993, Host B. tabaci & T.
abutiloneus, On Lycopersicon cheesmanii,
R. S. Stauffer (11 2, 2 ¢ mounted in Hoy-
er’s). Paratypes deposited with BMNH,
MJR, TAMU, UCR, and USNM.
Other specimens examined.—USA: Tex-
as: Bexar Co., San Antonio, San Antonio
Botanical Garden greenhouse, 11.vii.1991,
Bemisia tabaci on Hibiscus lasiocarpus, C.
Moomaw (1 2); Brazos Co., College Sta-
tion, Crop Biosciences Bldg. GH, TAMU,
18—22.xi1.1993, Bemisia tabaci on tomato,
RS. stautter (20), 2): Collese4Station;
TAMU culture, 7.x.1994, Bemisia tabaci on
Hibiscus, M. Hunter (14 2, 7 3); College
Station, TAMU Biotech Greenhouse,
7.x.1994, Bemisia on tomato, M. Hunter (1
3); College Station, TAMU BC Labs,
411.1995, Bemisia on Hibiscus, M. Hunter
(1 3d). All specimens from MIR.
Eretmocerus tejanus Rose and
Zolnerowich, new species
(Figs. 26-28, 33)
Diagnosis.—Females of E. tejanus can
be distinguished by the club that is 6.2—
7.1X as long as wide (Fig. 26), pedicel that
is 3—3.7X as long as wide and 0.29-0.36X
as long as the club, mesoscutum with 6 se-
tae, and gastral tergite II usually with 2
pairs, but occasionally 1 pair, of lateral se-
tae (Fig. 28).
Females of E. tejanus are very similar to
E. corni and E. eremicus. Females of E.
corni have the pedicel 2.4—3.1X as long as
wide and 0.25—0.30X as long as the club
(Fig. 6), and have not been reliably record-
ed from Bemisia (tabaci complex). Eret-
mocerus eremicus has the anterior alary
fringe of the forewing slightly longer (Fig.
11), and gastral tergite II usually has 1 pair,
but occasionally 2 pairs, of lateral setae
(Fig. 12).
Males of E. tejanus can be distinguished
by the longitudinal fuscous bands on the
mesocutum (Fig. 33). Males of E. corni
lack longitudinal fuscous bands on the me-
socutum, and males of E. eremicus have the
mesoscutum fuscous along the anterior
margin (Fig. 30), or with a light fuscous
“‘T’? shape (as in Fig. 31), or evenly fus-
cous.
Female.—Length of critical point-dried
specimens 0.44—0.59 mm. Head and body
varying from white testaceous to tan yel-
low, with darker specimens having the ver-
tex orange yellow. Eyes grey red, ocelli
clear. Legs testaceous, tarsi occasionally
slightly darker than tibiae. Wings hyaline.
Specimens mounted in Hoyer’s with head
amber and body pale yellow.
Face and occiput with transverse substri-
gulate sculpture, interscrobal area vertically
substrigulate. Antenna (Fig. 26) with radi-
VOLUME 99, NUMBER 1
Figs. 26-28.
cle 4.2 as long as wide; scape 5X as long
as wide, 2.1 X length of radicle, 1.6 length
of pedicel, 0.6 length of club; pedicel 3—
3.7X as long as wide, 1.4 length of rad-
icle; 0.6X length of scape, 0.29—-0.36x
length of club. Funicle I triangular, 1.1 as
wide as long; funicle II] subquadrate. Club
6.2—7.1X as long as wide, 1.8 length of
scape, 2.8 length of pedicel. Antennal ra-
toy 1 2.2:14:3 5.
Mesoscutum trapezoidal and with 6 se-
tae, anterior % with reticulate sculpture,
substrigulate sculpture in narrow triangular
medial area, remainder with elongate retic-
ulate sculpture. Parapsis with 2 setae, with
fine reticulate sculpture proximally and sub-
strigulate sculpture distally; axilla with |
seta and substrigulate sculpture. Scutellum
with 4 setae and 2 placoid sensilla lateral
to and closer to posterior setae, and elon-
gate substrigulate sculpture. Propodeum
with faint substrigulate sculpture medially
Eretmocerus tejanus. 26, 2 antenna. 27, 2 forewing. 28, 2 gaster.
and around spiracles. Endophragma extend-
ing into gastral tergite II.
Forewing (Fig. 27) 2.9 as long as wide
at width I, 2.6 as long as maximum width
of disc. Longest anterior alary fringe 0.13 <
width of disc, longest posterior alary fringe
0.28 width of disc. Base of wing usually
with | seta, occasionally 1—3, distal portion
of costal cell usually with 2—4, occasionally
1—4, setae. Marginal vein with 3 long setae,
9-12 setae, occasionally 9-14, between
marginal vein and linea calva. Linea calva
closed posteriorly by setae, with 10—15 tu-
bercles on ventral surface of wing near pos-
terior end of linea calva; a group of 26—37
setae including those forming distal edge of
linea calva point toward anterior margin of
wing, remaining 162—208 setae in disc
point to distal apex of wing. Ratio of sub-
marginal:marginal:stigmal veins 3:1.3:1.
Hind wing 7.3 as long as wide and with
3—8 setae in the center.
24 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 29-32. do habitus. 29, Eretmocerus debachi. 30, E. eremicus. 31, E. furuhashii. 32, E. staufferi.
Gastral tergite I with substrigulate sculp-
ture, smooth around lateral setae, lateral
margins faintly imbricate with stippling;
gastral tergites I-VI (Fig. 28) usually with
paired setae as follows: 1, 2, 2, 2, 2, 1, al-
though specimens are known with 1-2, O-
3, 2, 2-3, 2-3, 1. Syntergum with 4 setae.
Ovipositor slightly exserted, equal in
length to club, 2.8X length of pedicel, 1.7
length of scape, equal in length to midtibia.
VOLUME 99, NUMBER 1
Fig. 33. Eretmocerus tejanus, 3 habitus.
Male.—Length of critical point-dried
specimens 0.43—0.51 mm. Critical point-
dried specimens colored as female except
pronotum brown dorsolaterally, mesoscu-
tum with 2 sublateral longitudinal fuscous
bands, pale medially and laterally (Fig. 33),
scutellum slightly fuscous, foretarsi slightly
darker than tibia.
Specimens mounted in Hoyer’s with head
amber. Radicle and scape fuscous, with
scape darker at apex; pedicel dark fuscous;
multiporous plate sensilla of club fuscous.
Pronotum fuscous. Mesoscutum dark fus-
cous along anterior margin, slightly less
fuscous along length of mesoscutum, me-
dially and laterally pale, creating 2 longi-
tudinal fuscous bands (Fig. 33). Parapsis
and axilla slightly fuscous. Scutellum fus-
cous to slightly fuscous, usually darker
along anteror and posterior margins. Meta-
notum slightly fuscous, darker medially.
Propodeum fuscous except medially and
along anterolateral margins. Gaster pale to
slightly fuscous. Aedeagus brown. Foreleg
except coxa fuscous; midleg except coxa
pale, midtibia occasionally fuscous, midtar-
si brown; hind trochanter pale to slightly
fuscous, hind femur slightly fuscous, tibia
es
33
darker, tarsi brown. Base of forewing and
costal cell slightly fuscous; submarginal
vein brown fuscous, marginal and stigmal
veins slightly fuscous. Venation of hind
wing brown fuscous.
Host.—Reared from Bemisia (tabaci
complex).
Discussion.—Eretmocerus tejanus has
been referred to as E. californicus and E.
sp. nr. californicus. Although females of E.
tejanus are extremely similar to those of E.
eremicus, differences in male pigment pat-
terns, in conjunction with reciprocal mating
tests and electrophoretic analysis between
populations from Arizona and California,
and Texas (Hunter et al. 1996), indicate that
the Texas population forms a species dis-
tinct from the populations in Arizona and
California.
This species was one of the most com-
mon naturally occurring parasites of Bemi-
sia (tabaci complex) in the Rio Grande Val-
ley of south Texas prior to the 1995 Boll
Weevil Eradication Program. Eretmocerus
tejanus was not recovered from rearing
samples taken during this program. Popu-
lations of this species began to appear again
during the spring of 1996, approximately
26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
nine months following cessation of insec-
ticide treatments. Sampling for this and oth-
er species of parasites in the Rio Grande
Valley continues.
Eretmocerus tejanus has been released in
central and southern California for control
of Bemisia (tabaci complex).
Etymology.—From Spanish Tejano,
meaning ““Texan’’.
Holotype.—? mounted in balsam on a
slide labelled, “‘USA: Texas/Hidalgo Co./
Mission/Biological/Control/Laboratory/F,
lab culture/M94003”’ ‘‘original/material:/
USA: Texas/Hidalgo Co./Mission/18.i.
1994”’ “J. Rodriguez/ex: Bemisia/tabaci/
on: cabbage’’. Deposited in USNM.
Paratypes.—USA: same data as holotype
(3 2 and 2 6 mounted in balsam, 14 @ and
10 36 card-mounted); Texas: Hidalgo Co.,
Weslaco, TAES greenhouse, xi.1990, Host
Bemisia tabaci, On melon, B. Breene (6 2,
4 $6 mounted in Hoyer’s); Weslaco,
10.vi.1993, Host Bemisia sp., On Brassica
oleraceae [sic], C. Moomaw (19 @, 15 ¢
mounted in Hoyer’s). Paratypes deposited
with BMNH, MJR, TAMU, UCR, and
USNM.
Other specimens examined.—Texas:
Brazos Co., College Station, TAMU, B.C.
garden, (23*v-1991-" Bemisia tabacr, C.
Moomaw (1 @, 1 2); TAMU, B.C. garden,
3.vii.1991, Bemisia tabaci on kale, C.
Moomaw (1 6d); TAMU Green House,
6.vi.1991, Bemisia tabaci on squash, C.
Moomaw (7 @, 8 ¢); Hidalgo Co., Hargill,
25.vi.1993, Bemisia tabaci on cabbage,
Moomaw (12 @, 13 ¢); Hargill, 25.vi.1993,
Bemisia tabaci on kenaf, Moomaw (14 &,
6 3); nr. Hargill, Rio Farms, 2.viii.1993,
Bemisia tabaci on kenaf, Moomaw (8 2, 5
3); nr. Mission, Holbrook, 2.vi.1993, Be-
misia tabaci on watermelon, Moomaw (3
2,5 3); nr. Mission, Holbrook, 29.vi.1993,
Bemisia tabaci on kale, Moomaw (2 @); nr.
Mission, Holbrook farms, 26.vii.1993, Be-
misia tabaci on pigweed, C. Moomaw (10
2, 8 3); Weslaco, 9.vii.1991, Bemisia ta-
baci on cotton, M. Rose (4 9, 1 3); Wes-
laco, 25.i11.1992, B. tabaci on cantaloupe,
Moowaw (1 <6); Weslaco, USDA-ARS,
26.i11.1992, B. tabaci on sunflower, Moo-
waw (1 2, 1 3); Weslaco, USDA-ARS,
26.ii1.1992, B. tabaci on sunflower, Woolley
(1. 2, I ¢); Weslaco, USDA-ARS,
18.vii.1992, Bemisia tabaci on kale,
Moomaw (5 ¢, 6 3); Weslaco, USDA-ARS
test plot refuge, 18.vii.1992, Bemisia tabaci
on kale, M. Rose & C. Moomaw (6 &, 3
3); Weslaco, USDA-ARS, 30.x.1992, Be-
misia tabaci on kale, Moomaw (13 ¢,9 3);
Weslaco, Rio Grande Valley, 1.iv.1993, Be-
misia tabaci on cabbage, M. Rose (2 9, 1
3); Weslaco, Rio Grande Valley, 1.iv.1993,
Bemisia tabaci on collards, M. Rose (4 @&,
7 3); Weslaco, TAES, 20.iv.1993, Bemisia
tabaci on canola, Moomaw (3 2, 1 3@);
Weslaco, TAES-Riley, 28.vi.1993, Bemisia
tabaci on sunflower, C. Moomaw (5 &, 5
3); Weslaco, TAES-Riley, 27.vii.1993, Be-
misia tabaci on sunflower, Moomaw (12 @,
4 6); Weslaco, TAES-Riley, 31.vii.1993,
Bemisia tabaci on kale, Moomaw (9 @, 9
3); Weslaco, TAES, 21.viii.1993, Bemisia
tabaci on cotton, Moomaw (3 2, 5 3);
Weslaco, Rio Grande Valley, 19.v.1994, Be-
misia tabaci on mixed host plants, M. Rose
(25 2, 20 3). All specimens from MUIR.
ACKNOWLEDGMENTS
This research was supported by funds
from USDA/APHIS, USDA/ARS, USDA/
National Research Initiative, the Southern
Regional Sustainable Agriculture and Edu-
cation Program, and the Texas Higher Edu-
cation Coordinating Board. The work of G.
Zolnerowich was supported by a USDA/
APHIS/National Biological Control Institute
Postdoctoral Fellowship in Systematics.
We express our appreciation to M. E.
Schauff (Systematic Entomology Labora-
tory, USDA) and J. M. Heraty (UCR) for
their review of the manuscript and loan of
types. We thank FE D. Bennett and G. Evans
(University of Florida), C. Pickett and J.
Ball (CDFA), K. Hoelmer, J. Goolsby, and
L. Wendel (USDA/APHIS), R. Carruthers
and G. Butler (USDA/ARS), and D. Head-
rick (UCR) for their kind collaboration and
VOLUME 99, NUMBER I
for providing specimens. We thank Ben
Shaw for the drawings.
LITERATURE CITED
Brown, J. K. and J. Bird. 1992. Whitefly-transmitted
geminiviruses and associated disorders in the
Americas and the Caribbean Basin. Plant Disease
76: 220-225.
Brown, J. K., D. R. Frohlich, and R. C. Rosell. 1995.
The sweetpotato or silverleaf whiteflies: biotypes
of Bemisia tabaci or a species complex? Annual
Review of Entomology 40: 511—534.
Byrne, D. N., T. S. Bellows, Jr, and M. P. Parrella.
1990. Whiteflies in agricultural systems, pp. 227—
261. In Gerling, D., ed., Whiteflies: Their Bio-
nomics, Pest Status, and Management. Intercept,
Ltd. 348 pp.
Compere, H. C. 1936. Notes on the classification of
the Aphelinidae with descriptions of new species.
University of California Publications in Entomol-
ogy 6: 277-321.
Costa, H. S., M. W. Johnson, D. E. Ullman, A. D.
Omer, and B. E. Tabashnik. 1993. Sweetpotato
whitefly (Homoptera: Aleyrodidae): analysis of
biotypes and distribution in Hawaii. Environmen-
tal Entomology 22: 16—20.
Dozier, H. L. 1932. The identity of certain whitefly
parasites of the genus Eretmocerus Hald., with de-
scriptions of new species (Hymenoptera: Aphel-
inidae). Proceedings of the Entomological Society
of Washington 34: 112-118.
Ebeling, W. 1957. La mosca blanca lanuda de los ci-
tricos. Direccion General de Agricultura, Divul-
gaciones e informaciones No. 18, 9 pp.
Gill, R. J. 1992. A review of the sweetpotato whitefly
in southern California. Pan-Pacific Entomologist
68: 144-152.
Haldeman, S. S. 1850. On four new species of He-
miptera of the genera Ploiaria, Chermes, and
Aleurodes, and two new Hymenoptera, parasitic
in the last named genus. American Journal of Sci-
ence 9: 108-111.
Headrick, D. H., T. S. Bellows, Jr., and T. M. Perring.
1995. Behaviors of female Eretmocerus sp. nr.
californicus (Hymenoptera: Aphelinidae) attack-
ing Bemisia argentifolii (Homoptera: Aleyrodi-
dae) on sweet potato. Environmental Entomology
24: 412-422.
Headrick, D. H., T. S. Bellows, Jr., and T. M. Perring.
1996. Behaviors of female Eretmocerus sp. nr.
californicus (Hymenoptera: Aphelinidae) attack-
ing Bemisia argentifolii (Homoptera: Aleyrodi-
dae) on cotton, Gossypium hirsutum, (Malvaceae)
and melon, Cucumis melo (Cucurbitaceae). Bio-
logical Control 6: 64—75.
Howard, L. O. 1895. Revision of the Aphelininae of
North America. United States Department of Ag-
riculture Technical Series No. 1, 44 py
Howard, L. O. 1908. On two new species of parasites
of Aleyrodidae. Proceedings of the Entomological
Society of Washington 10: 63—65.
Hunter, M. S., M. FE Antolin, and M. Rose. 1996.
Courtship behavior, reproductive relationships,
and allozyme patterns of three North American
populations of Eretmocerus nr. californicus (Hy-
menoptera: Aphelinidae) parasitizing the whitefly
Bemisia sp., tabaci complex (Homoptera: Aley-
rodidae). Proceedings of the Entomological Soci-
ety of Washington 98: 126—137.
Mound, L. A. and S. H. Halsey. 1978. Whitefly of the
World. British Museum (Natural History) and
John Wiley and Sons. 340 pp.
Polaszek, A., G. A. Evans, and FE D. Bennett. 1992.
Encarsia parasitoids of Bemisia tabaci (Hyme-
noptera: Aphelinidae, Homoptera: Aleyrodidae): a
preliminary guide to identification. Bulletin of En-
tomological Research 82: 375-392.
Rose, M. 1988. Twenty years of biological control
success on Citrus in the United States, pp. 1153-—
1161. Jn Goren R. and K. Mendel, eds., Proceed-
ings of the Sixth International Citrus Congress,
Tel Aviv, Israel. Philadelphia: Balaban Publishers.
Rose, M. and D. Rosen. 1991-1992. Eretmocerus de-
bachi n. sp. (Hymenoptera: Aphelinidae), an ef-
fective parasite of Parabemisia myricae (Homop-
tera: Aleyrodidae). Israel Journal of Entomology
25-26: 199-207.
Rose, M. and G. Zolnerowich. 1994. Eretmocerus fu-
ruhashii n. sp. (Hymenoptera: Aphelinidae), an ef-
fective parasite of Parabemisia myricae (Kuwana)
(Homoptera: Aleyrodidae) in Japan. Japanese
Journal of Entomology 62: 285-292.
Rose, M. and P. DeBach. 1991-1992. Biological con-
trol of Parabemisia myricae (Kuwana) (Homop-
tera: Aleyrodidae) in California. Israel Journal of
Entomology 25: 73-95.
Rose, M., G. Zolnerowich, and M. Hunter. 1996. Sys-
tematics, Eretmocerus, and Biological Control,
pp. 477-497. In Gerling, D., ed., Bemisia 1995:
Taxonomy, biology, damage, control and manage-
ment. Intercept, LTD. 702 pp.
Rosen, D. and P. DeBach. 1979. Species of Aphytis
of the World (Hymenoptera: Aphelinidae). Dr. W.
Junk BV. 801 pp.
Schauff, M. E., G. A. Evans, and J. M. Heraty. 1996.
A pictorial guide to the species of Encarsia (Hy-
menoptera: Aphelinidae) parasitic on whiteflies
(Homoptera: Aleyrodidae) in North America. Pro-
ceedings of the Entomological Society of Wash-
ington. 98: 1—35.
Viggiani, G. and G. Evans. 1992. Descriptions of
three new species of Amitus Haldeman (Hyme-
noptera: Platygasteridae), parasitoids of known
whiteflies from the New World. Bolletino del La-
boratorio di Entomologia Agraria “Filippo Silves-
tri’ di Portici 49: 189-194.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 28-36
A REVIEW OF THE SHORE-FLY GENUS DIPHUIA CRESSON
(DIPTERA: EPHYDRIDAE)
WAYNE N. MATHIS
Department of Entomology, MRC 169, Smithsonian Institution, Washington, DC 20560,
U.S.A.
Abstract.—Diphuia Cresson is reviewed to include (1) the synonymy of D. nasalis
Wirth with D. nitida Sturtevant and Wheeler, (2) the description of D. flinti, new species
(Dominican Republic: El Seibo), and (3) a cladistic analysis of the genus.
Key Words:
While conducting field work on the Do-
minican Republic in 1995, a tiny, somewhat
shiny, black shore fly of the genus Diphuia
Cresson was discovered that is apparently
associated with freshwater habitats. Other
species of Diphuia occur along shorelines
and beaches of maritime environments with
brackish-water or saline habitats. Determin-
ing the identity of this species, which is less
than 2.5 mm in length, has led to this re-
view and cladistic analysis of Diphuia.
Diphuia is known only from the New
World, mainly from subtropical and tropical
zones, and has greatest diversity in the Ca-
ribbean and adjacent areas. Although com-
paratively common in nature, specimens are
scarce in collections, probably because of
the restricted distribution of the genus and
the small size of specimens (2.5 mm or
less). The nomenclatural history of Diphuia
and included species, as well as other avail-
able information were included in my ear-
lier revision (Mathis 1990) and are not re-
peated here except for the taxa being treat-
ed:
METHODS
The terminology and methods used in
this study were explained previously (Math-
is 1990). Because of the small size of spec-
Diptera, Ephydridae, shore flies, Diphuia, New World tropics, cladistics
imens, study and illustration of the male
terminalia required the use of a compound
microscope. To better assure effective com-
munication about structures of the male ter-
minalia, I have adopted the terminology of
other workers in Ephydridae (see references
in Mathis 1986). Usage of these terms,
however, should not be taken as an endorse-
ment of them from a theoretical or morpho-
logical view over alternatives that have
been proposed (Griffiths 1972, McAlpine
1981). Rather, I am deferring to tradition
until the morphological issues are better re-
solved.
Two venational ratios are used common-
ly in the descriptions and are defined here
for the convenience of the user (ratios are
averages of three specimens).
1. Costal vein ratio: the straight line dis-
tance between the apices of veins R,,, and
R,,;/distance between the apices of veins
R, and R,.,.
2. M vein ratio: the straight line distance
along M between crossveins (dm-cu and
r-m)/distance apicad of crossvein dm-cu.
The specimens used in this study are pri-
marily in the National Museum of Natural
History (USNM), Smithsonian Institution.
A few, especially primary types, are in the
American Museum of Natural History
VOLUME 99, NUMBER 1
(AMNH), New York. I also examined spec-
imens from the University of Texas, Austin
(UTA) collection.
The phylogenetic analysis was performed
with the assistance of Hennig86 ©, a com-
puterized algorithm that produces clado-
grams on the basis of parsimony. Before
performing the analysis, the character data
were arranged in transformation series and
then polarized primarily using outgroup
procedures.
Diphuia Cresson
Diphuia Cresson, 1944:4. Type species: Di-
phuia anomala Cresson, 1944, by origi-
nal designation; 1946:138, 140 [note,
key ].—Sturtevant and Wheeler, 1954:248
[notes].—Wirth, 1956:4 [discussion of
species]; 1965:737 [Nearctic catalog];
1968:5 [Neotropical catalog].—Mathis,
1990:746-756 [revision].—Mathis and
Zatwarnicki, 1995:155—156 [world cata-
log].
Description.—Mostly black, subshiny to
shiny, microtomentum usually sparse; small
shore flies, length 1.35—2.10 mm. Head:
Wider than high; face width-to-head width
ratio 0.28; frons black, mostly unicolorous,
lacking distinctively colored ocellar trian-
gle; frons wider than long, frontal length-
to-width ratio 0.58; frontal vestiture vari-
able; ocellar seta well developed, inserted
slightly in front of alignment of anterior
ocellus and at about the same distance apart
as between posterior ocelli; pseudopost-
ocellar setae usually well developed, length
subequal to ocellar setae, proclinate, slight-
ly divergent; 1 reclinate and 1 proclinate
fronto-orbital seta present, reclinate seta in-
serted slightly anteromediad of proclinate
seta; both inner and outer vertical setae
present; ocelli arranged to form isosceles
triangle, with distance between posterior
ocelli larger than between anterior ocellus
and either posterior ocellus. Antenna exert-
ed; pedicel with well-developed, proclinate,
dorsal seta; aristal length subequal to anten-
nal length and bearing 4—5 dorsal rays, with
~
basal 3 rays longer than apical 1—2, the lat-
ter subequal. Eye apparently bare of micro-
setulae (using a stereomicroscope). Face
black in both sexes and with silvery white,
microtomentose antennal grooves and with
2 lines, sometimes irregular, paralleling
parafacials, these and similarly invested and
colored ventral margin (microtomentum
sometimes interrupted at middle) form a fa-
cial triangle that has a small microtomen-
tose area below facial prominence; face not
carinate between antennal bases but slight-
ly, conically protrudent at middle (best seen
in lateral view); ventral facial margin shal-
lowly emarginate; face bearing 2 facial se-
tae, the dorsal seta very slightly larger, both
inserted near parafacials; parafacials dense-
ly microtomentose, silvery white; clypeus
very sparsely microtomentose, black; pal-
pus blackish brown to black; mouthparts
not geniculate, labella shorter than medi-
proboscis. Thorax: Generally black, vesti-
ture of microtomentum variable with spe-
cies, although generally sparse; pleural ar-
eas lacking stripes of distinctly colored mi-
crotomentum. Chaetotaxy with mesonotal
setae poorly developed except for those at
posterior margin; mesonotal setulae numer-
ous and not arranged in well-defined setal
tracks; prescutellar acrostichal setae much
larger than other acrostichal setulae and
more widely set apart; only 1 dorsocentral
seta, inserted posteriad; intra-alar setulae ir-
regularly seriated; presutural seta well de-
veloped, length subequal to notopleural se-
tae; 2 scutellar setae and scutellar disc with
sparse, scattered setulae; postpronotal seta
1; postalar seta 1; notopleural setae 2, in-
sertion of posterior seta elevated dorsally
above anterior one; anepisternal setae 2, in-
serted along posterior margin; katepisternal
seta well developed, conspicuous. Wing:
membrane mostly hyaline to very slightly
milky white; veins behind costa pale, usu-
ally yellowish to yellowish brown; vein
R,,, extended well beyond level of cross-
vein dm-cu, 2nd costal section at least 1%
times longer than 3rd section; alular mar-
ginal setulae short, less than % alular height.
30 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Legs: femora black; tibiae dark basally,
concolorous with femora, apices yellowish.
Abdomen: Fifth segment of male well scler-
otized, elongate, not normally visible from
a dorsal view, usually retracted within 4th
segment; Sth tergum and sternum of male
united anteriorly to form a complete annu-
lus. Male terminalia as follows: cercus rod
shaped, bearing 1—3 conspicuously longer
setae at ventral margin; surstylus well de-
veloped, well sclerotized, conspicuous, and
as long or longer than cercus.
Distribution. —New World. Temperate to
tropical zones, in North America along the
east coast (New York south to Florida) and
the Caribbean to Colombia and Ecuador in
South America.
Diagnosis.—Diphuia can be distin-
guished from other genera of Hecamedini
as follows (characters indicated by an as-
terisk (*) are autapomorphies that corrobo-
rate the monophyly of Diphuia): *colora-
tion very dark, usually black; *microtomen-
tum of head and thorax generally sparse,
giving a subshiny to faintly dull appear-
ance; facial coloration of male and females
similar, lacking sexual dimorphism; face,
although slightly protrudent medially (best
seen in profile), not acutely pointed in lat-
eral view; *face with silvery white micro-
tomentose markings, antennal grooves, 2
vertical lines, ventral margin, an area below
the facial prominence, and parafacials; pre-
sutural and prescutellar setae well devel-
oped; *pleural region lacking a stripe or
stripes; Sth segment of male well sclero-
tized and its tergum moderately elongate;
*5th tergum and sternum of male united an-
teriorly to form a complete annulus; and
male genitalia with distinct, well-sclero-
tized, elongate surstyli.
KEY TO SPECIES GROUPS AND
SPECIES OF DIPHUIA
1. Anepisternum with anteroventral %4—% bare,
shiny black, otherwise with sparse investment
of whitish gray microtomentum; mesonotum
sparsely invested with microtomentum, subshi-
ny; Sth tergum of male with anterior margin
even (Fig. 2); surstylus with ventral margin
broadly rounded; gonite trapezoidal; aedeagus
Walle Ayes < lONNNIRERMIES Ale SSS Bos 55555005
ae irhs ricci Ue D. nitida Sturtevant and Wheeler
— Anepisternum almost entirely invested with
whitish gray microtomentum; mesonotum
moderately densely microtomentose, golden
brown; 5th tergum of male shallowly or deeply
emarginate (Fig. 16); surstylus truncate ven-
trally, pointed apically; gonite with basal half
subrectangular; aedeagus not bifurcate apically
nN
. Ventral portion of face with median, vertical,
densely microtomentose, silvery white stripe
extended dorsad from and connected to micro-
tomentose ventral margin; 5th tergum longer
than wide, anterior margin shallowly emargin-
Fl (teeneons muo ee A ls lord ohare oe D. flinti, new species
— Ventral portion of face with median, densely
microtomentose, silvery to golden white spot
(frequently triangular or diamond shaped) that
is not directly connected with microtomentum
along ventral margin; 5th tergum wider than
long, anterior margin deeply emarginate, emar-
gination V-shaped
3. Surstyli long and narrow, length subequal to
that of cercus; gonite pointed posteroventrally;
aedeagus only moderately curved apically .. .
Pte the ene eae eats Menke D. anomala Cresson
— Surstyli moderately short and robust, length
shorter than cercus; gonite with posteroventral
portion broadly bifurcate; aedeagus more
curved apically, point oriented anteriorly
roe teria2 tate ciated Oy Gav D. zatwarnickii Mathis
Diphuia nitida Sturtevant and Wheeler
(Figs. 1-8)
Diphuia nitida Sturtevant and Wheeler,
1954:248.—Wirth, 1965:737 [Nearctic
catalog].—Mathis, 1990:754 [revi-
sion]|.—Mathis and Zatwarnicki, 1995:
156 [world catalog].
Diphuia nasalis Wirth, 1956:3; 1968:5
[Neotropical catalog].—Mathis, 1990:
750-754 [revision].—Mathis and Za-
twarnicki, 1995:156 [world catalog].
New Synonym.
Description.-Small shore flies, length
1.35 to 1.75 mm.
Head: Frons sparsely invested with fine
brownish microtomentum, microtomentum
becoming sparser or lacking on 2 small ar-
eas laterad of posterior ocelli, 2 spots along
the anterior margin, and sometimes a small
VOLUME 99, NUMBER 1
spot in front of the anterior ocellus. Ventral
portion of face with median, densely micro-
tomentose, silvery to golden white spot
(frequently triangular or diamond shaped)
that is not directly connected with micro-
tomentum along ventral margin.
Thorax: Mesonotum sparsely invested
with fine brownish to golden brown micro-
tomentum, mostly subshiny to shiny; ane-
pisternum with anteroventral 4% to 2 bare of
microtomentum, shiny, posterodorsal por-
tion invested with fine, grayish microto-
mentum. Wing with costal vein ratio 0.58;
M vein ratio 0.40.
Abdomen: 5th tergum with anterior mar-
gin essentially straight, at most very shal-
lowly arched anteriorly (Fig. 2); 5th ster-
num undivided, as a narrow band connected
dorsally with anteroventral portion of 5th
tergum (Fig. 1). Male terminalia (Figs. 3—
8) as follows: epandrium narrow in lateral
view, much higher than wide (Fig. 3); sur-
stylus as long as cercus but almost twice its
width, broadly rounded apically (Fig. 3);
gonite in lateral view parallelogram-shaped,
posterior angles produced into pointed pro-
cesses, posteroventral process sinuous
(Figs. 4-5); aedeagal apodeme rounded an-
teroventrally (Figs. 4—5); aedeagus acutely
pointed apically, in dorsal or ventral view
bifurcate apically (Figs. 4—5); hypandrium
in ventral view wider than long, anterior
margin shallowly arched anteriorly (Figs. 5,
7-8).
Type material—The holotype 2 of D.
nitida 1s labeled *‘Doug]l[a]s[t]on[,] L[ong].
I[sland]., Nfew]. Y[ork].[,] Au[gust]. 17,
[19]52/HOLOTYPE Diphuia nitida Stvt &
Whlr [pink]/TYPE 6695 [dark pink; num-
ber handwritten].’”” The holotype is point
mounted, is in poor condition (the antennae
and several setae are missing and the left
side of the body and wings are covered par-
tially with glue), and is deposited in the
ANSP (6695). Sturtevant and Wheeler stat-
ed that this specimen is a male, but it is
clearly a female.
The holotype 2 of D. nasalis is labeled
“Long Island[,] Deadman’s Cay[,] March
ai
11, 1953/Van Voast-A.M.N.H. Bahama Isls.
Exped Coll. E.B.Hayden/d/3 HOLOTYPE
Diphuia nasalis W.W. Wirth [red, gender and
species name handwritten].’’ The holotype
is point mounted, is in good condition (tip
of right wing folded back on itself), and is
deposited in the AMNH. Although the ho-
lotype was listed as a male (Wirth 1956:4)
and the specimen is so marked, it is a fe-
male.
Other specimens examined.—BAHA-
MAS. Crooked Island, Landrail Point, 5
Mar 1953, E. B. Hayden, L. Giovannoli (1
2; AMNH); Exuma Cays, Staniard Bay, 13
Jan 1953, E. B. Hayden (1 2; AMNH);
Long Island, Deadman’s Cay, 11 Mar 1953,
E. B. Hayden (2 ¢6; AMNH, USNM). BE-
LIZE. Stann Creek District: Bread and But-
ter Cay, 25 Mar 1988, W. N. Mathis (5 6,
1 2; USNM); Glover’s Reef (Long Cay,
Middle Cay, Northeast Cay, Southwest
Cay), 26-28 Jul 1989, W.N. Mathis (29 6,
12 2; USNM); Man of War Cay, 8-15 Nov
1987, W. N. & D. Mathis (7 6, 4 @;
USNM); Twin Cays (West Bay), 22 Mar
1988, W. N. Mathis (1 6, 1 9; USNM);
Wee Wee Cay, 24-25 Mar 1988, 21 Jul
1989, W. N. Mathis (5 ¢; USNM). Six Be-
lize, 1959, N. L. H. Krauss (1 6; USNM).
BERMUDA. Hamilton Parish. Shelly Bay,
20 Nov 1987, D. J. Hilburn, N. E. Woodley
(2 2; USNM). DOMINICAN REPUBLIC.
Barahona: Cabral (canals E of Cabral;
18°152'N,. 71°13:4 W); 16; May 1995: W.
N. Mathis (1 ¢, 1 2; USNM). La Romana:
Isla Saona, Catuano (18°11.7'N, 68°46.8’W),
13 May 1995, W. N. Mathis (1 ¢; USNM).
GRAND CAYMAN. Governor Gore Bird
Sanctuary (19°16.7'N, 81°18.5’W), 25 Apr
1994, W. N. Mathis (1 6; USNM). JA-
MAICA. Falmouth (bay shore), 1 Mar 1969,
W. W. Wirth (1 d6; USNM); Milk River
Bath (mangroves), 11 Mar 1970, T. Farr, W.
W. Wirth (5 3, 1 2; USNM); Negril Beach
(rocky shore), 12 Mar 1970, W. W. Wirth
(1 6; USNM); Runaway Bay (bay shore),
16-28 Feb 1969, W. W. Wirth (3 o;
USNM). UNITED STATES. Florida. Mon-
roe Co., Bahia Honda Key (seashore), 11
Go
N
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-8.
terminalia (epandrium, cercus, surstyli), lateral view. 4, Gonite, lateral view. 5, Internal male terminalia (gonite,
hypandrium, aedeagal apodeme, aedeagus), lateral view. 6, Aedeagus and aedeagal apodeme, lateral view. 7,
Hypandrium, aedeagal apodeme, and aedeagus, ventral view. 8, Hypandrium, lateral view. Scale = 0.1 mm.
Apr 1970, W. W. Wirth (4 36, 3 2; USNM);
Big Pine Key, 11-Apr-30 Dec, 1954, 1970,
HV Weems (i'd, 1 2; USNM): Vong
Key, 23) Jum 1953; Mi’R> Wheeler (lc:
UTA); Saddlebunch Keys, 29 Dec 1953, H.
V. Weems (1 ¢, 1 2; USNM). North Car-
olina. Onalow Co., Ashe Island, 11 Aug
1975, J Cy Dukes “(1"2>-USNM)-
Diphuia nitida. 1, Sth tergum and sternum, lateral view. 2, 5th tergum, dorsal view. 3, Male
Distribution.—Nearctic: USA (FL, NC,
NY). Neotropical: Bahamas, Belize, Ber-
muda, West Indies (Dominican Republic,
Grand Cayman, Jamaica).
Natural history—The vast majority of
specimens from the Belizean cays were col-
lected by sweeping just above mangrove
peat that is well shaded most of the day. A
VOLUME 99, NUMBER |
few specimens, apparently feeding, were
collected on flowers. The association with
mangrove peat must be opportunistic, as the
species occurs in areas where mangrove
does not now exist.
Diagnosis.—This species is distinguished
from congeners, especially D. anomala and
D. zatwarnickii, by the sparsely microto-
mentose mesofrons; the subshiny mesono-
tum that is very sparsely invested with fine,
golden brown microtomentum; the sparsely,
whitish gray microtomentose anepisternum;
the shiny anterior half of the katepisternum;
and several characters of the male termi-
nalia (see description and figures).
Remarks.—Since 1989, when the re-
search from my earlier revision (Mathis
1990) was completed, I have studied longer
series of specimens from additional locali-
ties. These studies indicate that D. nasalis
and D. nitida are conspecific, with the latter
being the senior synonym. Although no
more specimens from the type locality
(New York: Long Island, Douglaston) of D.
nitida have become available, I now strong-
ly suspect that the difference noted earlier
between the nominate species has mostly to
do with the poor condition of the holotype
female of D. nitida. This specimen, which
is the entirety of the type series, appears to
have been “‘rubbed”’ (antennae and several
setae are missing), which could also ac-
count for the shinier frons, the only distin-
guishing character.
Diphuia flinti Mathis, new species
(Figs. 9-16)
Description.—Small to moderately small
shore flies, length 1.60 to 2.10 mm.
Head: Frons sparsely and uniformly in-
vested with fine brownish to grayish micro-
tomentum. Ventral portion of face with me-
dian, vertical, densely microtomentose, sil-
very white stripe extended dorsad from and
connected to microtomentose ventral mar-
gin.
Thorax: Mesonotum, including postpro-
notum and notopleuron, sparsely invested
33
with fine brownish to golden brown micro-
tomentum, mostly subshiny; anepisternum,
anepimeron, and posterior half of katepi-
sternum sparsely invested with fine, whitish
gray microtomentum; anterior half of kat-
episternum with oval spot bare of micro-
tomentum, shiny. Wing with costal vein ra-
tio 0.58; M vein ratio 0.40.
Abdomen: 5th tergum with anterior mar-
gin emarginate medially (Fig. 16); Sth ster-
num a narrow arch that is connected ante-
riorly with 5th tergum (Fig. 11), apex with
2 flanges (Fig. 12). Male terminalia (Figs.
9—10, 13-15) as follows: epandrium higher
than wide in lateral view, ventral margin
pointed (Fig. 10); surstylus longer than cer-
cus, obtusely angulate in lateral view, apex
truncate and pointed medially and anteri-
orly (Figs. 9-10); gonite in lateral view
with basal half subrectangular, with narrow,
sinuous, pointed process apically (Figs. 13—
14); aedeagal apodeme in lateral view ir-
regularly triangular (Fig. 13); aedeagus
rounded apically in lateral view (Fig. 13),
pointed (not bifurcate) in dorsal view (Fig.
14); hypandrium in ventral view wider than
long, anterior margin deeply arched with
medial margins recurved, posterior margin
evenly arched except for a short, median
process (Fig. 15).
Type material——The holotype ¢ is la-
beled ““DOMINICAN REPUB. El Seibo:
near Rincon[,] 18°45.3'N, 68°55.7’W{[,]
12 May 1995, WNMathis/USNM ENT
00136995 [bar code label].”’ The allotype
2 and 14 paratypes (10 6, 4 2) bear the
same locality label as the holotype. Other
paratypes are as follows: DOMINICAN RE-
PUBLIC. “EV Setbo> EV Seibo"G@* kim E;
18°44.73'N, 68°59.2'W; 120 m), 12 May
1995, W.N. Mathis (1 ¢,5 2; USNM). The
holotype is double mounted (minuten in sil-
icon plastic block), is in excellent condi-
tion, and is deposited in the USNM.
Distribution.—Neotropical. Dominican
Republic (El Seibo).
Diagnosis.—I can distinguish this species
only on the basis of structures of the male
terminalia, especially the shape of the 5th
34 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
SORES
CEES
S37
rhe
S
vA
9
Figs. 9-16. Diphuia flinti. 9, Epandrium, cerci, and surstyli, posterior view. 10, Same, lateral view. 11, 5th
tergum and sternum, lateral view. 12, Apex of Sth sternum, ventral view. 13. Gonite, aedeagus, and aedeagal
apodeme, lateral view. 14, Left gonite and aedeagus, dorsal view. 15, Hypandrium, ventral view. 16, 5th tergum,
dorsal view. Scale = 0.1 mm.
tergum, surstylus, gonite, aedeagus, and hy-
pandrium as described and illustrated.
Natural history.—The type series was
collected along the banks of the Rio Qui-
sibani, which is a freshwater stream.
Etymology.—It is a pleasure to name this
species after Dr. Oliver S. Flint, Jr, who
accompanied me on the field work that re-
sulted in the collection of this species.
PHYLOGENETIC CONSIDERATIONS
Diphuia is in the tribe Hecamedini
(Mathis 1991a, 1991b, 1993) and belongs
to the clade including Hecamede Haliday
and Allotrichoma Becker, sensu lato, in-
cluding Pseudohecamede Hendel (Mathis
1991la). This relationship is evidenced by
the retracted fifth tergum of the male, which
is moderately elongate, almost tubular. Di-
phuia may be the sister group to the rest of
the Allotrichoma clade, following Cresson’s
(1944) original assessment, although the
evidence supporting this relationship is not
compelling, and Diphuia could be mostly
closely related to Hecamede Haliday.
The monophyly of Diphuia within He-
camedini is well established (see generic di-
agnosis for a partial list of synapomor-
phies), and the remainder of this section
will be devoted to the phylogenetic rela-
tionships among the species. First the char-
acter evidence is presented, followed by a
brief discussion that is intended to comple-
ment the cladogram.
Character evidence.—The characters
used in the cladistic analysis are listed be-
VOLUME 99, NUMBER 1
Table 1.
analysis of Diphuia.
Matrix of characters used in the cladistic
Characters
Taxa 1 2 Sy 4.5 6 7 8 9
Outgroug ORO OO. tO) 105 OR SOR 0)
D. anomala 1 0 1 2 | | 1 i] 1
D. zatwarnickii 1 (0) 1 2 1 1 1 1 1
D. nitida 0 1 0 0 10) 10) 0 0 0
D. flinti Oe 05. 10 1 1 1 1 1 0
low and the distribution of their states is
shown in the matrix (Table 1) and discussed
here. For each character, the plesiomorphic
condition is indicated by a O, and the rela-
tive apomorphic condition(s) is indicated by
al or 2.
—
. Vestiture of mesonotum: (0) sparsely mi-
crotomentose, subshiny; (1) moderately
densely microtomentose.
. Vestiture of anepisternum: (0) sparsely,
uniformly microtomentose; (1) antero-
ventral % to % bare, shiny.
. Length of 5th tergum of male: (0) longer
than wide (Figs. 1—2, 11, 16); (1) wider
than long (Figs. 1—2, 16—17 in Mathis
1990).
. Shape of anterior margin of 5th tergum
of male: (O) straight, even; (1) shallowly
emarginate; (2) deeply emarginate,
V-shaped (Figs. 2, 17 in Mathis 1990).
Shape of ventral margin of epandrium:
(O) broadly formed, somewhat truncate;
(1) tapered to point.
Shape of surstylus: (0) rounded; (1)
truncate ventrally, pointed.
Shape of gonite: (0) trapezoidal with
sides more or less equal (Fig. 4); (1)
somewhat rectangular, longer than high
(Bias 13):
. Shape of posterior margin of hypan-
drium: (0) smooth and even; (1) with a
small, median projection.
Nw
eS)
&
fo)
SN
=a
(oe)
9. Shape of aedeagus: (0) slender, 3—4>
longer than high in lateral view; (
more robust, length in lateral view about
twice height (Figs. 7, 21 in Mathis
1990).
1)
Discussion.—Analysis of the character
evidence (Table 2), all morphological and
primarily from the male terminalia, resulted
in a single cladogram (Fig. 17). The single
tree resulted from the implicit enumeration
(ie*) option of Hennig86 and has a length
of 10 steps and consistency and retention
indices of 1.00 and 1.00, respectively. The
analysis of characters is given in Table 2.
The cladogram (Fig. 17) for the four spe-
cies consists of three bifurcations, with D.
nitida at the base and the remaining species
as the sister group. The monophyly of the
latter assemblage is established by charac-
ters four (1) and five through eight. The
next bifurcation has D. flinti, which 1s
known thus far only from the Dominican
Republic, branching off and as the sister
group to D. anomala and D. zatwarnickii.
The monophyly of the D. anomala and D.
zatwarnickii group is indicated by four
characters (1, 3;:-4(2), 9):
ACKNOWLEDGMENTS
I am grateful for the assistance in the
field from Oliver S. Flint, Jr. (USNM) and
Kelvin A. Guerrero (Museo Nacional de
Historia Natural, Santo Domingo). For the
opportunity to examine specimens housed
in their collections, I thank David Grimaldi
and Julian Stark (AMNH) and Donald Azu-
ma (ANSP). For critically reviewing a draft
of this paper, I thank Allen L. Norrbom.
The illustrations were skillfully inked by
Elaine R.S. Hodges.
Funding for this research project, es-
Table 2. Analysis of characters used in the cladistic analysis.
Character: 1 2 3
Steps: 1 2
Consistency Index: 100 100 100.
Retention Index: 100 100 100
4 5 6 if 8 9
| 1 1 1 1 |
100 100. 100 100 100 100
100 100 100 100 100 100
36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
outgroup
(2)
Fig. 17.
consistency index 1.00, retention index 1.00.
pecially the field work in Belize and the
Dominican Republic, was provided by the
Caribbean Coral Reef Ecosystems (CCRE)
and Biodiversity Program (Biological Sur-
veys and Inventories, BSI), National Mu-
seum of Natural History, Smithsonian In-
stitution. This is contribution number 501
of the CCRE project, which is partially sup-
ported by a grant from the Exxon Corpo-
ration.
LITERATURE CITED
Cresson, E. T., Jr. 1944. Descriptions of new genera
and species of the dipterous family Ephydridae.
Paper XIV. Notulae Naturae. The Academy of
Natural Sciences of Philadelphia 135: 1-9.
. 1946. A systematic annotated arrangement of
the genera and species of the Neotropical Ephyd-
ridae (Diptera). 1. The subfamily Psilopinae.
Transactions of the American Entomological So-
ciety 71: 129-163.
Griffiths, G. C. D. 1972. The Phylogenetic Classifi-
cation of Diptera Cyclorrhapha with Special Ref-
erence to the Structure of the Male Postabdomen.
Series Entomologica 8: 1-340. W. Junk, Hague.
Mathis, W. N. 1986. Studies of Psilopinae (Diptera:
Ephydridae), I: A revision of the shore fly genus
Placopsidella Kertész. Smithsonian Contributions
to Zoology 430: 1-30.
. 1990. A revision of the shore-fly genus Di-
phuia Cresson (Diptera: Ephydridae). Proceedings
of the Entomological Society of Washington
92(4): 746-756, 22 figures.
1991a. Studies of Gymnomyzinae (Diptera:
Ephydridae), Il: A revision of the shore-fly sub-
asaya
Cladogram depicting hypothetical relationships among species of Diphuia. Tree length 10 steps,
D. nitida
D. flinti
D. anomala
D. zatwarnickiil
genus Pseudohecamede Hendel of the genus AI-
lotrichoma Becker. Smithsonian Contributions to
Zoology 522: 28+iii pp.
. 1991b. Classification of the shore flies (Dip-
tera: Ephydridae), past, present, and future, pp.
209-227. In Weismann, L., et al., eds., Proceed-
ings of the Second International Congress of Dip-
terology. VEDA, Bratislava, SPB Academic Pub-
lishing, 368 pp. The Hague.
1993. Studies of Gymnomyzinae (Diptera:
Ephydridae), IV: A revision of the shore-fly genus
Hecamede Haliday. Smithsonian Contributions to
Zoology 541: 46+iu1 pp.
Mathis, W. N. and T. Zatwarnicki. 1995. A world cat-
alog of the shore flies (Diptera: Ephydridae).
Memoirs on Entomology, International 4: vi+423
Pp.
McAlpine, J. E 1981. Morphology and terminology-
adults, pp. 9-63. In McAlpine, J. F, et al., eds.,
Manual of Nearctic Diptera. Ottawa. [Volume | is
Monograph 27 of Research Branch Agriculture
Canada. |
Sturtevant, A. H. and M. R. Wheeler. 1954. Synopses
of Nearctic Ephydridae (Diptera). Transactions of
the American Entomological Society 79: 151—
PSM fe
Wirth, W. W. 1956. The Ephydridae (Diptera) of the
Bahama Islands. American Museum Novitates
1817: 1-20.
1965. Family Ephydridae, pp. 734-759. In
Stone, A., et al., eds., A catalog of the Diptera of
America north of Mexico. U.S. Department of Ag-
riculture, Agriculture Handbook 276.
1968. 77. Family Ephydridae, pp. 1—43. In
Papavero, N., ed., A catalogue of the Diptera of
the Americas south of the United States. Depar-
tamento de Zoologia, Secretaria de Agricultura,
Sao Paulo.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 37—41
A NEW SPECIES OF DRYMUS FIEBER FROM MEXICO, WITH A KEY TO
SPECIES AND A CHECKLIST OF WESTERN HEMISPHERE DRYMINI!
(HEMIPTERA: LYGAEIDAE)
JAMES A. SLATER AND HARRY BRAILOVSKY
(JAS) Department of Ecology and Evolutionary Biology, University of Connecticut,
Storrs, CT 06269, U.S.A; (HB) Departamento de Zoologia, Instituto de Biologia, UNAM,
Apartado Postal No. 70-153, Mexico 04510, D. FE, Mexico.
Abstract.—The first Mexican species of Drymus Fieber, is described as Drymus mex-
icanus, n. sp., from the mountains of the state of Veracruz. A key to the Western Hemi-
sphere species is included together with a list of the Western Hemisphere species of the
tribe Drymini. A discussion of the distribution of the tribe is given and the affinities of
the Nearctic fauna analyzed.
Key Words:
The zoogeography of the Rhyparochrom-
inae was reviewed in detail by Slater
(1986). In that paper he noted the interest-
ing distribution of the members of the tribe
Drymini and how closely the overall distri-
bution paralleled that of the tribe Rhypar-
ochromini. Both of these tribes are abun-
dant, diverse and speciose in the Palearctic,
Ethiopian-Oriental (= Paleotropical) and
Australian regions. However, both make up
only a very small proportion of the Nearctic
rhyparochromine fauna and both are absent
from the the true Neotropical Region (in-
cluding the West Indies). Of the Nearctic
fauna most species belong to genera that are
also found in the Palearctic.
The Drymini tend to be associated with
woodland margins and to live in moderately
mesic habitats. The Rhyparochromini occur
for the most part in relatively dry savannah-
like habitats.
The absence from the Neotropics of
Drymini, despite their abundance in Afri-
ca, suggests a radiation subsequent to 90
million years b.p. or subsequent to the
Drymus, Mexico, biogeography, distribution, checklist
breakup of West Gondwanaland. It also
suggests that despite the abundance of the
Drymini in Australia they are relatively re-
cent elements in that fauna since there was
apparently no movement of such a fauna
across Antarctica (and because there are no
native species in New Zealand). The most
plausible scenario for the occurrence of
Drymini in the Western Hemisphere ap-
pears to be as a Holarctic element that was
present during, or before, the onset of the
Pleistocene. If this is true, it means that
two taxa have been able to reach generic
status during this period (both of which are
confined to areas of California that escaped
glaciation).
The Mexican drymine fauna is found in
montane areas and all species are members
of genera also found in the United States
and Canada as well as in the Palearctic.
Drymus is the most unusual of these in that
the two previously known species are
known from only a few localities in the
western United States.
All measurements are in millimeters.
38 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 1.
Drymus mexicanus, Slater and
Brailovsky, new species
(Figs. 1, 2)
Robust, subelliptical. Head, anterior
pronotal lobe, scutellum and antennae dark
chocolate brown, almost black. Explanate
pronotal margins, posterior pronotal lobe,
entire hemelytron, legs and labium dull red-
dish-brown. Fourth antennal segment with
apical two-thirds white. Coarsely punctate
Over entire surface except wing membrane,
OR Re
~~
Sai
Sey iy,
Drymus mexicanus. Dorsal view, head and pronotum.
the punctures on head and anterior pronotal
lobe relatively small.
Head acuminate, non-declivent; tylus at-
taining distal one third of first antennal seg-
ment; vertex convex. Eyes set well away
from anterior margin of pronotum. Length
head 0.84, width 1.00, interocular space
0.82. Pronotal calli confluent or nearly so,
mesally, elevated above surface of posterior
lobe. Lateral pronotal margins broadly ex-
planate (Fig. 1); posterior margin of pro-
notum shallowly concave. Length prono-
VOLUME 99, NUMBER I
Fig. 2. Drymus mexicanus. Abdomen, lateral view.
tum 1.20, width 2.00. Scutellum with a me-
dian elevation distally, mesally concave on
basal half. Length scutellum 1.12, width
1.20. Hemelytra with lateral margins broad-
ly convex. Clavus with four rows of punc-
tures. Length claval commissure 0.60. Mid-
line distance apex clavus-apex corium 1.24.
Midline distance apex corium-apex mem-
brane 0.64. Metathoracic scent gland auri-
cle curved posteriorly. Evaporative area
only slightly exceeding end of auricle, dis-
tally truncate, covering only mesal one-
third to one-fourth of metapleuron. Fore fe-
mur only moderately incrassate, armed be-
low on distal one fourth with a short sharp
spine with three minute spines distad. La-
bium extending to metacoxae, first segment
not attaining base of head. Length labial
segments I 0.62, II 0.70, III 0.54, [IV 0.44
(approx.). Length antennal segments I 0.50,
II 0.80, TT 0.62, [V 0.68. Total body length
5:65.
All abdominal spiracles located below
sternal shelf. Posterior pair of trichobothria
on sternum four located dorsoventrally, the
ventral trichobothrium reduced and obso-
lete (Fig: 2):
Holotype 2 MEXICO: Veracruz: Na-
39
olinco, 1.VIHI.1977, 2100 m. In Universi-
dad Nacional Autonoma de Mexico collec-
tion.
This is a very distinctive species at once
separable from the other Western Hemi-
sphere species by the white distal portion
of the fourth antennal segment, the much
more strongly explanate lateral pronotal
margins, and the shorter antennae. Actually
this is not the only species of Drymus with
a pale distal end to the fourth antennal seg-
ment; several Palearctic species have a
somewhat paler apex on segment four, al-
though none that we have examined have a
white terminal end to the antennae.
KEY TO WESTERN HEMISPHERE
SPECIES OF DRYMUS
1. Distal half of fourth antennal segment white,
strongly contrasting with dark coloration of ba-
sal half of antennae; explanate margins of pro-
notum broad, subequal to width of second an-
tennal segment; second antennal segment sub-
equal in length to length of head
Ee Cenc ee Ia eer et oot ae en mexicanus N. sp.
— Fourth antennal segment unicolorous dark
brown to black; explanate lateral pronotal mar-
gins relatively narrow, much narrower than
width of second antennal segment; second an-
40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tennal segment considerably longer than length
Ofte ad! Oe spe Fee os hee) aon die eee 72
. Large (6.5—7 mm); very dark almost black; an-
terior and posterior pronotal lobes nearly even-
ly punctate; expanded lateral margins of pro-
notum concolorous with doraal surface of pro-
NOUN seh Se EG oes eee crassus Van Duzee
— Smaller (rarely over 5 mm); dull brown; ante-
rior pronotal lobe more finely punctate than
posterior lobe; expanded lateral margins of
pronotum usually slightly paler than surface of
PLONOtUM: «5 0% Ges. ges eenate siete unus (Say)
i)
CHECKLIST OF WESTERN HEMISPHERE
DRYMINI
See Slater 1964 and Slater and O’Don-
nell 1995 for complete references.
Drymus Fieber
crassus Van Duzee 1910. New England
south to Florida, west to Texas and South
Dakota.
mexicanus, n. sp. Mexico.
unus (Say) 1831. Eastern Canada, south to
North Carolina, west to Colorado.
Eremocoris Fieber
arnaudi Brailovsky 1982. Mexico.
borealis (Dallas) 1852. Western U.S.A.
canadensis Walley 1929. British Columbia;
Idaho.
chalmaensis Brailovsky and Barrera 1981.
Mexico.
cupressicola Ashlock 1979. California.
depressus Barber 1928. New England south
to Florida and Louisiana coastal areas.
dimidiatus Van Duzee 1921. Colorado.
extremus Brailovsky and Cervantes 1989.
Mexico.
ferus (Say) 1832. Widespread over much of
U.S.A. and southern Canada.
garciai Brailovsky and Barrera 1981. Mex-
ico.
guerrerensis Brailovsky and Barrera 1981.
Mexico.
inquilinus Van Duzee 1914. California.
legionarius Brailovsky and Barrera 1981.
Mexico.
lopezformenti Brailovsky and Barrera 1981.
Mexico.
melanotus Walley 1929. British Columbia;
Idaho.
mimbresianus Brailovsky and Cervantes
1989. Mexico.
obscurus Van Duzee 1906. California; Ida-
ho; British Columbia.
opacus Van Duzee 1921. California.
semicinctus Van Duzee 1921. California;
Idaho.
setosus Blatchley 1926. Eastern U.S.A.
squalidus Brailovsky and Barrera 1981.
Mexico.
Gastrodes Westwood
arizonensis Usinger 1938. Arizona.
conicolus Usinger 1933. California.
intermedius Usinger 1938. British Colum-
bia.
pacificus (Provancher) 1886. Nebraska and
Colorado, west to Calfornia and British
Columbia.
walleyi Usinger 1938. Ontario, British Co-
lumbia.
Scolopostethus Fieber
atlanticus Horvath 1893. Northern U.S.A.
and southern Canada; Newfoundland to
British Columbia south to New Jersey
and New Mexico.
diffidens Horvath 1893. Northern U.S.A.
and southern Canada: Nova Scotia to
British Columbia south to New Jersey.
pacificus Barber 1918. California, Idaho,
British Columbia.
thomsoni Reuter 1874. Almost throughout
Holarctic.
tropicus (Distant) 1882. California to Gua-
temala.
Thylochromus Barber
nitidulus Barber 1928. California.
Togodolentus Barber
wrighti (Van Duzee) 1914. California.
ACKNOWLEDGMENTS
We express our appreciation to Mr. Er-
nesto Barrera (UNAM) Mexico for prepa-
ration of the illustrations and to Dr. Jane
VOLUME 99, NUMBER 1
O’Donnell, University of Connecticut, for
assistance with the manuscript.
LITERATURE CITED
Slater, J. A. 1964. A Catalogue of the Lygaeidae of
the World. 2 Volumes. University of Connecticut.
Storrs. 1688 pp.
4]
. 1986. A synopsis of the Zoogeography of the
Rhyparochrominae (Heteropteraa: L)
Journal of the New York Entomological Society
94: 262-280.
Slater, J. A. and J. E. O'Donnell. 1995. A Catalogue
of the Lygaeidae of the World (1960-1994). New
York Entomological Society, 410 pp.
gaeidae).
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 42—49
NESTING BEHAVIOR OF KROMBEINICTUS NORDENAE LECLERCQ,
A SPHECID WASP WITH VEGETARIAN LARVAE
(HYMENOPTERA: SPHECIDAE: CRABRONINAE)
KARL V. KROMBEIN AND BETH B. NORDEN
Department of Entomology, National Museum of Natural History, Smithsonian Insti-
tution, MRC 165, Washington, DC 20560, U.S.A.
Abstract.—Nesting behavior of the recently described Sri Lankan wasp, Krombeinictus
nordenae Leclercq, is discussed. Females nest in the hollow internodes of the leguminous
myrmecophyte, Humboldtia laurifolia Vah\. The biology of this stem-nesting crabronine
iS unique among Sphecidae in several aspects. An adult female exhibits remarkable ma-
ternal care, rearing one larva at a time, and feeding it progressively. Progressive provi-
sioning has not been noted previously for any Crabroninae. Nests lack cell partitions and
mature larvae are transported to the basal regions of their stems for cocoon spinning. The
cocoon is also unlike that of any other known crabronine species, exhibiting adaptations
to internode morphology and allowing movement of adults within the nest cavity. Finally,
K. nordenae is remarkably different from all other known Sphecidae in feeding pollen
rather than paralyzed arthropod prey to its larvae.
Key Words:
cocoon, pollen
Sri Lanka has been called the land of ser-
endipity. A recent example is the amazing
nesting behavior of a Ceylonese wasp, a
newly described genus and species, Krom-
beinictus nordenae Leclercq (1996). This
pretty little wasp (Fig. 1), 5—6 mm long, has
creamy to pale yellow markings on its black
head and thorax, and a mostly light red ab-
domen bearing narrow, transverse, brown to
black stripes on some of the dorsal seg-
ments. Krombeinictus belongs to the Sphe-
cidae, normally a family of predaceous,
mostly solitary wasps.
MATERIALS AND METHODS
Our first encounter with this species was
when we received a single male of K. nor-
denae from a colleague, Prof. Fred R. Rick-
son. It was among a few wasps that
Sphecidae, Crabroninae, Humboldtia, Sri Lanka, internode, myrmecophyte,
emerged from a dozen internodes of the
myrmecophyte, Humboldtia laurifolia Vahl,
that he collected in the Sinharaja Forest Re-
serve in Sri Lanka in 1992.
We visited Sri Lanka in 1993, and spent
five days (18-20 Jul and 2—3 Aug) in the
rainforest near Gilimale, Ratnapura District,
06°46'N, 80°26’E. We hoped to make be-
havioral observations on Krombeinictus
during this brief period, but intermittent
rains of the delayed monsoon season pre-
cluded nesting activity by the wasps. How-
ever, we censused about a thousand inter-
nodes from H. laurifolia, and placed several
hundred unopened stems directly into al-
cohol for subsequent study. These inter-
nodes were split open carefully in the lab-
oratory to avoid damaging associated or-
ganisms, and notes were made on their con-
tents.
VOLUME 99, NUMBER |
43
Figs. 1-2.
Humboldtia laurifolia Vahl
(Figs. 3-6, 12—15)
This botanical section is condensed from
a detailed account of the plant’s morphol-
ogy in Krombein et al. (in prep.). Hum-
boldtia laurifolia (Fabaceae) (Fig. 3) occurs
only in Sri Lanka; three other species of the
genus and one variety occur in southern In-
dia. Humboldtia laurifolia is a small under-
story tree, growing to a height of about 10
m, found in the lowland rainforest of the
southwestern quadrant of Sri Lanka. It usu-
ally grows along streams or in seeps, is
highly gregarious, and occurs in groups of
ten to several hundred trees.
Humboldtia laurifolia and two of its In-
dian congeners, H. brunonis Wallich and H.
decurrens Beddome ex Oliver, are of par-
ticular interest to naturalists because they
are myrmecophytes that have coevolved
with several species of ants. The trees pro-
vide swollen internodes, each with a self-
opening entrance, that serve as domatia for
the ants. The trees also provide an abun-
dance of extrafloral nectaries on leaves,
stipules and inflorescences whose secre-
tions are attractive to the ants. In turn, the
ants protect the foliage, especially the
tender, young leaves, from herbivory.
Humboldtia laurifolia is typical of leg-
umes in having pinnately compound leaves,
each leaf having four to six pairs of oppo-
Krombeinictus nordenae, female. 1, Lateral view. 2, Frontal view of head.
site leaflets (Fig. 4). As in many legumes
there is a developmental period during
which a flush of four or five new inter-
nodes, each with an associated leaf, is pro-
duced over a period of two months. As the
internode develops, the apical part becomes
inflated, and is filled with pith. When the
internode matures, the pith collapses
against the inner wall, forming a hollow
cavity, and a slit-like opening develops near
the apex (Fig. 5). The opening widens grad-
ually until ants or other small organisms
can access the hollow domatium. Some oc-
cupants apparently gnaw at the more or less
elliptical opening, transforming it to an oval
or circular opening that is gradually rimmed
by a callus (Fig. 6), the plant’s response to
the injury. Aculeate Hymenoptera (ants,
wasps, bees) remove the pith to make the
cavity larger, except for one species of sol-
itary crabronine wasp, Crossocerus muka-
lanae Leclercq which uses the pith in con-
structing its nest in the cavity.
We found aculeates nesting in internodes
ranging in length from 4.3 to 11.2 cm. The
basal end of the internode is woody to a
variable extent, so the length of the cavity
varies from 3.2—6.3 cm. The woody outer
wall of the internode is 0.3—0.7 mm thick.
The internode entrances are elliptical to cir-
cular in shape with width to length mea-
surements ranging from a minimum of 1.1
x 1.1 mm to a maximum of 1.2 X 1.8 mm.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
hes
‘ os
dd
Figs. 3-6. Humboldtia laurifolia. 3, Habitat; collecting on plant. 4, Leaf; note swollen internode at lower
right. 5, Internode opening early in development. 6, Internode opening to Krombeinictus nordenae nest; note
callus around hole.
VOLUME 99, NUMBER I
Krombeinictus nordenae Leclercq
(Figs. 1, 2, 7-11)
Apparently an uncommon wasp, we
found only eight specimens (6 @, 2 d) in
Humboldtia internodes at Gilimale as com-
pared to several hundred of its fellow cra-
bronine, Crossocerus mukalanae. It has un-
usual morphology in that it is the only ge-
nus of Crabroninae, other than the Oriental
Vechtia Pate, in which both sexes possess
a triangular lamella overhanging the deep
scapal basin (Fig. 2).
The female is unusual morphologically in
the Crabroninae in lacking a pygidium de-
limited by carinae on the last abdominal ter-
gum. Instead there is a median brush of sev-
eral rows of close-set, erect setae (Fig. 7).
The last abdominal segment of females usu-
ally bears the residue of a secretion from
abdominal glands (Fig. 8) that we believe
may function as an ant guard. We suspect
that a nesting female uses its brush of setae
to smear this secretion around the nest en-
trance to deter predators and parasites from
entering while the adult wasp is foraging.
The Oriental Piyumoides Leclercq, con-
sidered by him (1996) to be the genus most
closely related to Krombeinictus, also lacks
a pygidium. Females, however, lack the me-
dian brush of setae on the last abdominal
tergum. There is no secretion from abdom-
inal glands on this segment in females of
three of the four known species in our col-
lection.
Nest.—We found only ten nests of K.
nordenae compared with about 75 of C.
mukalanae, and adult females were present
in only four of them. We believe that the
missing females took flight during the pe-
riod between gathering Humboldtia branch-
es, and placing the internodes in alcohol.
Several females were found sheltering in
empty internodes.
One typical nest exhibited the sequence
of nesting activity (Fig. 9). The internode
was 6.3 cm long, and the maximum width
was 4.0 mm. The interior cavity was 5.0 cm
long and had a maximum width of 3.4 mm.
45
When we split the internode, we found a
small wasp larva, 2 mm long, on the wall
of the cavity just below the entrance hole
The female was lower in the cavity, prob-
ably a reaction to being immersed in alco-
hol when the internode was _ preserved.
There was a wasp pupa with well developed
adult coloration in its cocoon at the bottom
of the cavity. Ten mm above this cocoon
was a second cocoon that contained a post-
defecated larva just prior to pupation.
From these data, and observations in oth-
er internodes with nesting K. nordenae, we
deduce the following behavioral sequence.
The foundress lays the first egg on the inner
cavity wall just below the entrance. When
the larva hatches, she feeds it progressively
until mature. The wasp then transports it to
the bottom of the cavity where it subse-
quently spins its cocoon, pupates, and slow-
ly begins to develop adult coloration. After
the wasp takes the mature larva below, she
lays a second egg just below the entrance.
That larva is fed progressively, and, when
mature, it is transported lower in the cavity
for cocoon spinning. The same cycle is re-
peated again with the wasp depositing an-
other egg below the entrance.
Typically, many sphecid wasps that nest
in pithy stems or borings in wood construct
nests containing a linear series of cells. The
cells are sealed by partitions that separate
siblings, thus preventing cannibalism. Fur-
ther, nests are normally closed before eggs
hatch so that females have no contact with
their progeny. Thus, the progressive feed-
ing, maternal attention, and lack of parti-
tions is noteworthy in K. nordenae.
Cocoon.—The cocoon also is unique
among the Crabroninae. The typical crab-
ronine cocoon is more or less ovoid, cir-
cular in cross section, the posterior end ta-
pers rather narrowly, and there is a pore at
the anterior end, as in Ectemnius paucima-
culatus (Packard) (Krombein, 1964). The
cocoon of Krombeinictus (Fig. 10) is broad-
ly ovoid, tapers very slightly posterad, and
lacks a pore at the anterior end. The upper
surface is only slightly convex so that in
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 7-11. Krombeinictus nordenae. 7, Female, apex of abdomen lateral oblique; note median brush of
setae on last tergum. 8, Female, apex of abdomen, lateral oblique from rear; note dried secretion on last segment,
and on fifth sternum. 9, Nest diagram. 10, Cocoon in section of internode from which adult emerged. 11, Cocoon
in section of internode, oblique; note ample space for passage of adults.
VOLUME 99, NUMBER 1
cross section the cocoon is more curved on
the side that is appressed against the round-
ed inner wall of the internode. Cocoon di-
mensions are 6—9 mm long, 2.6—3.4 mm
wide, and 2.0—2.1 mm high.
There is a space at least 2 mm high be-
tween its upper surface and the opposite in-
ner wall of the cavity (Fig. 11). A cocoon
of this shape permits the mother to crawl
over a cocoon to carry a mature larva to-
ward the lower end of the cavity, or for a
newly emerged adult from lower in the in-
ternode to crawl over a higher cocoon to
reach the nest entrance.
Larva.—The data above support our con-
clusion that the female feeds her larva pro-
gressively. We found larvae in various
stages of development, but never with any
prey or inedible prey fragments such as
wings and legs that one would expect to
find in the nest of a predaceous wasp. The
problem of larval food identity was finally
solved when we examined the exuviae of
post-defecated larvae, and adjacent fecal
wastes. Unmistakably, small grains of pol-
len had been excreted with the meconium
(Fig. 12) which were identical in size and
appearance with grains of Humboldtia pol-
len (Fig. 13). Also, we noted that freshly
preserved larvae had a distinct yellow color
reminiscent of that seen in bee larvae that
have fed on pollen. Variations in larval bee
color are attributed to pollen color (Norden,
1984).
Later in our investigation, Rickson sent
us a female of K. nordenae that he collected
from an internode in Gilimale. We found
clumps of Humboldtia pollen grains on the
mandibles and hypostomal setae beneath
the head (Figs. 14, 15). Due to the oily pol-
lenkitt that envelops the pollen grains, they
tend to adhere to each other and to insects
that come in contact with the stamens.
We infer from these data that the female
probably gathers a quantity of the clumped
Humboldtia pollen on the hypostomal area,
returns to the internode, and deposits the
pollen on the inner wall next to the head of
the larva. Again, this feeding behavior is
47
remarkable and in contrast to all known
sphecids whose larval food consists of par-
alyzed arthropods.
DISCUSSION
Behaviorally, K. nordenae is unusual or
unique among Sphecidae in several char-
acteristics of its life history. The female
manifests extraordinary maternal care, rear-
ing one larva at a time, and feeding it pro-
gressively. Progressive provisioning has not
been noted previously for any Crabroninae.
Evans (1966) reported that it developed in-
dependently at least four times in the Nys-
soninae. Evans also noted that eggs are pro-
duced more slowly in progressive provi-
sioners than in mass provisioners. However,
the slow development that characterized our
nests may actually be related to scarcity of
pollen and thus a slower rate of feeding
rather than to the rate of egg production.
Humboldtia flowering occurs throughout
the year, but the major production of inflo-
rescences is usually February to June.
The foundress with several brood in
varying stages of development forms a sub-
social group. Additional field observations
are needed to ascertain whether this sub-
social group reaches the level of social be-
havior by having a daughter join the foun-
dress in caring for later brood.
There is also the possibility that K. nor-
denae is so coadapted to Humboldtia that
the wasp nests only in the plant’s inter-
nodes, where nearby there is a plenitude of
extrafloral nectaries for adult feeding, and
pollen when the plant is in bloom. It is
noteworthy that the wasp was never col-
lected during the dozen years of the Smith-
sonian’s “Ceylon Insect Project,” 1968—
1980. None of the project specialists col-
lected on Humboldtia, but several of us, in-
cluding the first author on a number of trips,
collected at emergence holes in dead wood,
and in tunnels in wood, and failed to find
K. nordenae in these other plants. If K. nor-
denae is in fact associated only with Hum-
boldtia, we wonder if it or other species of
Krombeinictus may have a similar relation-
48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Seo) bee
Figs. 12-15. Humboldtia laurifolia pollen. 12, Pollen mass in meconium within cocoon of Krombeinictus
nordenae. 13, Grain from flower bud. 14, Pollen grains clumped on hypostomal setae behind mandibular apex
of female K. nordenae. 15, Pollen grains massed on setae beneath head of female K. nordenae.
ship in southern India with H. decurrens
and H. brunonis.
The cocoon also is unlike that of any oth-
er known crabronine. Clearly it is an ad-
aptation to the morphology of the internode
cavity and nesting behavior of the foun-
dress which permits movement of adults
throughout the cavity. The cocoon of the
normal twig-nesting crabronine is slightly
less in diameter than that of the cavity. The
larva spins a loose network of silken guy-
lines against the entire wall of the cavity
and then constructs the cocoon, suspending
it within the network (Krombein, 1964, fig
7b, shows cocoons not in contact with cell
wall). The cocoon of K. nordenae is spun
directly in contact with the rounded inner
wall of the internode, and there are only a
few silken guy-lines along the cocoon mar-
gins.
Finally, and perhaps most significant, the
species is extraordinarily different from all
other known Sphecidae in feeding its larvae
pollen rather than paralyzed arthropod prey.
VOLUME 99, NUMBER I
ACKNOWLEDGMENTS
We are grateful to our late colleague and
friend, PB. Karunaratne, for his many years
of help to scientists conducting field studies
in Sri Lanka. The Smithsonian Institution’s
“Ceylon Insect Project’? (1968-1980) was
highly successful in large part because of
Karu’s dedication and knowledge. During
our trip in 1993 he introduced the authors
to Humboldtia habitat, and helped collect
many of the internodes we examined. We
dedicate this work to him.
We also thank Arnold W. Norden who
was with us during part of our 1993 visit,
provided valuable support in field studies,
and reviewed an earlier draft of the manu-
script.
We gratefully acknowledge Prof. Fred R.
Rickson, Department of Botany and Plant
Pathology, Oregon State University, Cor-
vallis, for bringing H. laurifolia to our at-
tention, explaining the morphology and
phenology of the myrmecophyte, and for
insect specimens from his collecting in Sri
Lanka during 1992.
We are further indebted to Prof. Jean Le-
clercq, Liége, Belgium, for describing
Krombeinictus nordenae. It is an honor to
49
have our names bestowed on this bizarre
wasp with such unusual behavior
Within the Smithsonian we thank Susann
G. Braden who prepared most of the scan-
ning electron micrographs, and George L.
Venable who prepared the stem drawing
and photographic plates.
Finally, we thank Drs. Howard E. Evans,
Colorado State University, Fort Collins, and
Frank E. Kurczewski, State University of
New York, College of Environmental Sci-
ences and Forestry, Syracuse, for their re-
views and suggestions.
LITERATURE CITED
Evans, H. E. 1966. The comparative ethology and
evolution of the sand wasps. Harvard University
Press, Cambridge, Massachusetts. 526 pp.
Krombein, K. V. 1964. Natural history of Plummers
Island, Maryland, XVIII. The hibiscus wasp, an
abundant rarity, and its associates (Hymenoptera:
Sphecidae). Proceedings of the Biological Society
of Washington 77: 73-112.
Leclercq, J. 1996. A new genus and species of Cra-
bronini (Hymenoptera: Sphecidae) from Sri Lan-
ka. Memoirs of the Entomological Society of
Washington 17: 95-102.
Norden, B. B. 1984. Nesting biology of Anthophora
abrupta (Hymenoptera: Anthophoridae). Journal
of the Kansas Entomological Society 57: 243-
262.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 50-54
FENDEROMYIA SHAW, A VALID NORTH AMERICAN TAXON IN
MACROCERINAE (DIPTERA: MYCETOPHILOIDEA: KEROPLATIDAE)
Loic MATILE
Muséum national d’ Histoire naturelle, Laboratoire d’Entomologie et EP 90 du CNRS,
45, rue Buffon, F-75005 Paris, France.
Abstract.—The North American genus Fenderomyia Shaw, 1948, was long believed to
be a junior synonym of Macrocera Meigen, based on an artifact in thorax pleura and
several plesiomorphic states of characters. Identification of a specimen of the type species,
F. smithi Shaw, shows that the thoracic structure described by Shaw was not an artifact,
and makes it possible to point out several peculiar morphological characters of the taxon.
A phylogenetic analysis shows that these characters are autapomorphic and justifies the
resurrection of Fenderomyia. The genus is also present in the Neotropical Region.
Key Words:
analysis
The genus Fenderomyia was erected by
Shaw (1948) for a new North American
species, F. smithi, allied to Macrocera Mei-
gen, but which differed from this genus by
the following characters: base of median
vein distinct and reaching to the ““m-cu”’
crossvein, costa produced far beyond apex
of wing, and mesepimeron not reaching the
metapleuron.
Coher (1963) rightly pointed out that the
venation characters cited by Shaw for his
new genus occured also in several species
of Macrocera. He added that Shaw’s inter-
pretation of the pleural sclerites was erro-
neous, based on “‘a variable condition
shown by a series of M. brunnea Brunetti,
1912, to be a result of distortion during dry-
ing rather than of actual structure’. Ac-
cordingly, Coher considered Fenderomyia a
junior synonym of Macrocera.
Laffoon (1965) recognized Fenderomyia
as a valid genus, but it is doubtful that he
knew of Coher’s paper before the comple-
tion of his manuscript for the North Amer-
ican Catalog, which includes only ‘“‘some
Macrocerinae, Fenderomyia, valid generic status, morphology, character
1963 names’’. Thompson (1975), discuss-
ing the pleural morphology of Lygistorrhi-
na, mentions the synonymy established by
Coher, and Vockeroth (1981) does not in-
clude Fenderomyia in his key to the genera
of Nearctic Mycetophilidae, which means
that he also accepts Coher’ synonymy.
Matile (1990), revising the generic clas-
sification of Macrocerinae, discussed the
status of Fenderomyia and generally agreed
with Coher, noted that all three venation
characters were plesiomorphic for the
group, and accepted the thoracic distortion
explanation.
In some unsorted material of Macroceri-
nae at the National Museum of Natural His-
tory, Smithsonian Institution (USNM), I
identified a specimen belonging to Shaw’s
species. It was collected in Oregon (Coos
Co., Charleston, VI. 1963, NLH. Krauss),
as was Shaw’s type-series. As a result of an
examination of this specimen and compar-
ison with the extensive collection of Kero-
platidae available to me, it is now possible
to resurrect Fenderomyia from synonymy
VOLUME 99, NUMBER I
on the basis of several morphological char-
acters.
It is significant that the attribution to an
artifact of an exceptional feature, the
abridged mesepimeron, and the obviously
plesiomorphic venation characters cited by
Shaw for his new genus, have diverted the
attention of all subsequent authors, the pres-
ent one included, from the other very sig-
nificant characters cited in the original de-
scription.
MATERIAL AND METHODS
The USNM specimen is a male (accord-
ing to habitus) in rather poor condition: The
antennae, most of the legs, and the tip of
abdomen are broken (only the first five seg-
ments are present). It was originally glued
laterally to a piece of cardboard. The integ-
ument of the thorax was slightly transpar-
ent, thus showing the thoracic muscles; the
chitin of the pleura was smooth, without
trace of infolding or outfolding, and the
mesepimeron was definitely shortened. The
specimen was detached from the cardboard
and gently heated in distilled water, which
demonstrated that the ventral margin of the
mesepimeron corresponded to a true suture,
with an internal phragma. One of Shaw’s
diagnostic characters of Fenderomyia was
thus confirmed. The head was further boiled
in KOH to check the absence of a membra-
nous area between cerebral phragma and
front or occiput, thus confirming its posi-
tion in the Macrocerini rather than the Rob-
sonomyiini. The specimen was then dried
by the method of Sabrosky (1966), modi-
fied by the replacement of cellosolve by
monoethylene glycol (Matile 1994), and
then glued back to its cardboard. The head
was preserved in glycerine in a microvial.
According to the original description, the
type series of F. smithi should be in Shaw’s
collection, presently at the University of
Massachussetts. Dr. T.M. Peters was unable
to locate it either under Macrocera or Fen-
deromyia. Dr. R. J. Gagné was kind enough
to make inquiries about the possible loca-
tion of these specimens, but without suc-
cess, and writes “we will just hav
sume that the types were never distri
to the intended institutions and are probably
lost”’ (in litt. Apr. 1996).
~
-,
CHARACTER ANALYSIS
Four characters, presumably apomorphic,
can be added to the character of the short-
ened mesepimeron of Fenderomyia: the
long stem of the anterior fork; the diagonal
strip of dense, erect black setae of the met-
episternite, briefly mentioned in the original
description, which runs under the posterior
spiracle and above the dorsal margin of the
pleurite; the angular, short mediotergite,
strongly projecting backwards behind the
scutellum; and the presence of only one
spur on all tibiae. I have examined these
five features on about 100 species of Ma-
crocera, described or undescribed, from all
biogeographical regions, as well as in an
extensive collection of other Keroplatidae
and related families. The five characters are
discussed below.
1. Mesepimeron.—The evolution of the
mesepimeron has been discussed in Matile
(1990: 378, 411), and outgroup comparison
has shown that the most primitive condition
in most Nematocera was a vertical plate
reaching the lower margin of the pleura at
the level of the metepisternite. In the My-
cetophiloidea, the plate narrows ventrally,
and the disappearance of its ventral part is
undoubtedly apomorphic. The phenomenon
has occured several times in the Keroplati-
dae—at least twice in the Macrocerinae,
e.g. in Vockerothia Matile (Macrocerini)
and Micrepimera Matile and an unde-
scribed genus in Robsonomyiini—and once
in the Keroplatinae, namely in Nauarchia
Matile (Keroplatini). It is also a trait of all
Lygistorrhinidae and most Ditomyiidae.
2. Stem of anterior fork.—As regards the
stem of the media in the studied species of
Macrocera, the anterior fork always opens
far basad of the base of R;, except in two
Neotropical species, M. guayanasi Lane
and M. unidens Edwards. Among the other
Macrocerinae, a long stem as in Fendero-
52 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
myia is present only in a group of Austral-
asian Paramacrocera Edwards, and in the
Oriental genus Micrepimera Matile. In my
character analysis of the anterior fork of the
Mycetophiloidea (Matile 1990: 438), I not-
ed that fossil evidence was in favor of the
plesiomorphy of a long petiole, and inferred
that its shortness in the Keroplatidae was
apomorphic, and part of their groundplan.
The condition of Fenderomyia, Micrepi-
mera, the two South American Macrocera
and the Australasian Paramacrocera should
thus be a reversal, and its appearance in at
least four groups of Macrocerinae the result
of parallelism.
3. Postspiracular setae.—There is often
in Keroplatidae a sparse patch of short setae
at the posterior or ventral margin of the me-
tepisternite, but the dorsal rows of dense
setae in F. smithi do not really stand on the
metepisternite, but above it, on the peris-
piracular membrane, a few of the ventral
setae extending to the extreme dorsal mar-
gin of the sclerite (Figs la, b). These setae
should in fact properly be named infraspi-
racular setae. I have been unable to find this
feature in any Macrocerinae, in Arachno-
campinae and Keroplatinae (Keroplatini). I
have checked also representatives of almost
all the described genera of Orfeliini and
have found this character absent also. It
does not exist in the more primitive family
Ditomyiidae, nor in the presumed sister-
group of the Keroplatidae, the Diadocidi-
idae (for a provisional phylogenetic analy-
sis of the relationships of the families of
Mycetophiloidea, see Matile 1990: 383—
386). The apomorphic state of the character
seems therefore well founded.
4. Mediotergite.—In most Macrocera
known to me, the mediotergite is high, sub-
vertical in its upper half, then curves gently
downward and forward to meet the meta-
notum, thus not projecting behind the scu-
tellum, or very slightly so. In a few species,
the sclerite is evenly rounded and projects
somewhat behind the scutellum, the condi-
tion illustrated by Shaw for M. formosa
Loew, but never in such proportions as in
Fenderomyia (Fig. 1a; compare also Shaw’s
Figs. 1 and 2).
The evolution of the mediotergite has
been studied (Matile 1990: 405), and out-
group comparison has shown that its prim-
itive condition in Mycetophiloidea was a
vertical, high and slightly rounded sclerite.
In fact, this character is one of the nine used
to separate Macrocerini and Robsonomyi-
ini, the two tribes of Macrocerinae. Within
the subfamily, the projecting mediotergite
was considered autapomorphic for the Rob-
sonomyiini. The condition of Fenderomyia
thus must have appeared independently
once in Macrocerini.
5. Tibial spurs.—Only one tibia and three
tarsomeres, as well as some isolated tarso-
meres, remained on the piece of cardboard
of the USNM specimen and these became
detached as the glue dissolved; the absence
of a sensorial crypt indicates that the re-
maining tibia belongs to leg II or III; it has
only one very short spur. Shaw states “‘tibia
with one spur’’, and he had three specimens
available, therefore I assume that his obser-
vation refers also to both the median and
posterior legs.
The loss of the outer spurs II-III is an
evolutionary trend of the Keroplatidae (Ma-
tile 1990: 418) which occured indepen-
dently once in the Keroplatinae Keroplatini
(Xenokeroplatus Matile), and several times
in the Orfeliini, but the state of this char-
acter was not known up to now in the Mac-
rocerinae, although the reduction in size is
common. Most Macrocera known to me
have two posterior spurs, the longest about
twice as long as the width of the tibia at its
apex. Occasionally, the spurs are reduced to
one tibial width, or a little less. Only in M.
guayanasi and M. unidens are there two
very minute spurs as observed on the Ar-
gentinian Fenderomyia mentioned below.
DISCUSSION
The validity of Fenderomyia is supported
by five strong apomorphies. The short me-
dian fork is shared by two species of Mac-
rocera (both with a rather shortened medi-
VOLUME 99, NUMBER 1
AG)
lens, Al
and infraspiracular setae.
otergite), and does not exist elsewhere in
the subfamily. The thoracic structure has
appeared several times in the subfamily, but
never in Macrocera. The projecting, short-
ened mediotergite is unique for the Macro-
cerini, while it is characteristic of the Rob-
sonomyiini. The loss of the external tibial
spurs II-III is not yet known to happen in
the Macrocerinae. Lastly, the infraspiracu-
lar rows of setae are unique in the family
Keroplatidae and its closest relatives. This
amply justifies the acceptance of Fendero-
myia as a valid taxon, if only to emphasise
this peculiar set of apomorphies.
In the Duret Collection of Neotropical
Mycetophiloidea, now preserved in the Mu-
séum national d’ Histoire naturelle, Paris, I
found an unidentified female of “‘Macro-
cera” (Argentina, Salta, Campo Quijuano)
which shares with F. smithi the disappear-
ance of the ventral part of the mesepimeron,
the long stem of the median fork and the
projecting mediotergite, but lacks the dorsal
row of infraspiracular setae. Sc, is definitely
absent, but the basal fold of the media is
Nn
Las)
a b
Fenderomyia smithi (USNM specimen). a, Lateral view of thorax and coxae. b, Posterior spiracle
very faint. The two anterior legs and one
posterior leg are left; there is a short spur
on tibia I, and two very minute spurs on the
II, smaller than the basal diameter of the
protarsus. I am reluctant to describe for-
mally the species from a unique female, but
the specimen certainly represents the clos-
est known relative of F. smithi. Therefore,
the infraspiracular setae would be autapo-
morphic for F. smithi.
Provisionally, pending a revision of Ma-
crocera, | propose to follow Shaw and to
recognize Fenderomyia as a valid genus
close to Macrocera, with the following di-
agnosis:
Fenderomyia Shaw, 1948: 94. Type spe-
cies: Fenderomyia smithi Shaw, 1948: 94
(original designation).
A genus close to Macrocera, sharing
with this genus the globular antennal scape
and the long flagellum, but differing in the
incomplete mesepimeron, interrupted at the
level of lower third of the laterotergite, the
54 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
projecting mediotergite, the stem of the me-
dial fork long, ending at level of the base
of R;. In the type species, posterior spiracle
with several rows of posterior setae and
only one tibial spur II-III. In the Neotrop-
ical species, spiracular setae absent, and
hind tibia with two minute spurs.
Additional characters of taxonomic or
phylogenetic significance in Macrocera or
Macrocerini are: Cerebral sclerite large,
meeting the eyes at their anterior corner,
bearing long anterior setae. Thorax and
coxae as in Fig. la, acrostichal bristles ab-
sent. Scutellum with short, fine marginal
bristles. Mesanepisternite with long, erect,
dorsal setae. Subscutellar membranous area
narrow. Wing: C strongly produced after
R,, Sc, visible as a trace (USNM specimen)
or absent, basal fold of the media very
strong (F. smithi) or faint (Neotropical spe-
cies). Anal vein long, reaching to the mar-
gin. Wing membrane without macrotri-
chiae. Ciliation of veins, dorsal surface: C,
R,, branches of the anterior fork, M,, Cu,
and tip of anal vein. Male genitalia simple,
of the Macrocera type (Shaw, 1948: fig. 4).
Distribution: Nearctic and Neotropical.
ACKNOWLEDGMENTS
My thanks are due to Dr. Raymond J.
Gagné for the loan of an important material
of Keroplatidae from USNM, including the
specimen discussed above, and for his
search for the type series of Fenderomyia
smithi. Drs. R. J. Gagné and D. Williams
were kind enough to revise the language of
the manuscript. I thank also Gilbert Hode-
bert for the drawings.
LITERATURE CITED
Coher, E. I. 1963. Asian Macrocera Meigen, 1803,
(Diptera: Mycetophilidae), with some remarks on
the status of the genus and related genera. Bulletin
of the Brooklyn Entomological Society 58(1): 23—
36.
Laffoon, J. L. 1965. Family Mycetophilidae (Fungi-
voridae), pp. 196-229. In Stone, A. et al., eds., A
Catalog of the Diptera of America north of Mex-
ico. U.S. Department of Agriculture, Agricultural
Handbook 276, iv + 1—1696.
Matile, L. 1990. Recherches sur la systématique et
Vévolution des Keroplatidae (Diptera, Myceto-
philidae). Mémoires du Muséum national d’ His-
toire naturelle, sér. A, Zoologie 148: 1—682.
. 1994. Les Diptéres d’Europe occidentale. I.
Introduction, Techniques d’ étude et Morphologie.
Nématoceéres, Brachycéres Orthorrhaphes et As-
chizes. Paris, Boubée, 439 pp.
Sabrosky, C. W. 1966. Mounting Insects from Alco-
hol. Bulletin of the Entomological Society of
America 12(3): 349.
Shaw, E R. 1948. A new genus and species of fungus-
gnats (Mycetophilidae). Bulletin of the Brooklyn
Entomological Society 43(3): 94-96.
Thompson, FE C. 1975. Notes on the genus Lygistor-
rhina Skuse with the description of the first Ne-
arctic species (Diptera: Mycetophiloidea). Pro-
ceedings of the Entomological Society of Wash-
ington 77(4): 434-445.
Vockeroth, J. R. 1981. 14. Mycetophilidae, pp., 223—
246. In McAlpine, J. E, ed., Manual of Nearctic
Diptera. Volume 1.Biosystematics Researc Centre,
Research Branch. Agriculture Canada. Mono-
graph No. 28. Ottawa, vi + 674 pp.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 55-59
CLADISTICS AND BIOGEOGRAPHY OF THE ASSASSIN BUG GENUS
MELANOLESTES STAL (HETEROPTERA: REDUVITIDAE)
Maria del C. Coscar6n and Juan J. Morrone
(MCC) Departamento Cientifico de Entomologia, Museo de La Plata, Paseo del Bosque,
1900 La Plata, Argentina; (JJM) Laboratorio de Sistematica y Biologia Evolutiva (LAS-
BE), Museo de La Plata, Paseo del Bosque, 1900 La Plata, Argentina.
Abstract.—Melanolestes Stal is a monophyletic genus of nine species, two Nearctic
and seven Neotropical. A cladistic analysis of the genus was carried out using 28 char-
acters. The cladograms were rooted with the genus Peirates Serville. The analysis yielded
20 equally parsimonious cladograms, with 34 steps, CI = 0.52, and RI = 0.44. A suc-
cessive weighting procedure resulted in one cladogram with 77 steps, CI = 0.84, and RI
= (0.87. The distribution of Melanolestes coincides in part with a previous study on
Peiratinae, that showed that the former continuous Amazonian forest was separated into
two parts by a diagonal line of open areas. In addition, the two Nearctic species M.
picicornis and M. picipes are sister-taxa, so a single dispersal event accounts for the
presence of Melanolestes in the Nearctic.
Key Words:
The New World assassin bug genus Me-
lanolestes Stal (Heteroptera: Reduviidae:
Peiratinae) is known from southeastern
Canada to northern Argentina. The nine
species belonging to this genus have been
recently revised (Coscar6n and Carpintero
1994). Two of these species, M. picicornis
Stal and M. picipes (Herrich-Schaeffer), are
restricted to the Nearctic Region, whereas
the remainder are Neotropical. Within the
Neotropics, M. goiasensis Coscar6n and
Carpintero, M. lugens Coscar6n and Car-
pintero, M. minutus Coscar6n and Carpin-
tero, and M. picinus Stal have very small
distributional areas within the Amazonian
and Chacoan domains. In a previous bio-
geographic study (Morrone and Coscar6n
1996), we analyzed distributional patterns
of the Neotropical Peiratinae, concluding
that these patterns have been basically
caused by the gradual development of a di-
agonal line of open areas (Chaco-Cerrado-
Peiratinae, Reduviidae, cladistics, biogeography
Caatinga), which separated the former con-
tinuous tropical forest into two parts.
In this paper we provide a cladistic anal-
ysis of Melanolestes, and discuss its bio-
geographic patterns.
MATERIAL AND METHODS
This study is based on the revision of
Melanolestes by Coscar6n and Carpintero
(1994), and the examination of specimens
borrowed from the following collections:
American Museum of Natural History, New
York, USA; The Natural History Museum,
London, United Kingdom; Instituto Na-
cional de Pesquisas Amazonicas, Manaus,
Brazil; Museo Argentino de Ciencias Na-
turales Bernardino Rivadavia, Buenos Ai-
res, Argentina; Museo de La Plata, La Plata,
Argentina; Museu de Zoologia de Sao Pau-
lo, Sao Paulo, Brazil; Naturhistoriska Riks-
museet, Stockholm, Sweden; Zoologisches
Museum der Humboldt Universitat zu Ber-
56
character states; 1, 2 = apomorphic character states; ?
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Data matrix and characters of the species of Melanolestes used in the analysis. 0 = plesiomorphic
missing data.
No
ONAN ff WwW
\O
10.
lite
1
13).
14.
15).
16.
Ws
18.
19.
20.
2ile
D2r
23):
24.
25:
26.
PaTte
28.
Peirates OVDD0D0D0DD0D0D0D00D0D0000000000000000
M. argentinus CLONE OOM SOLO LO LOR OLO Ts eal ete OOKOLORIsO sia
M. degener AAO OS WO OOO POI eh OO) ea SL
M. goiasensis ORB ECOR ORR OLO MS eile sie Oy Oise rales?
M. lugens OZ MEO MEE ON 2 OMS Oss OM OFOTO OO Om eiiaG
M. minutus OZMFOLO MOA TSO ORO ME OLO;OLOl ON 25 cela?
M. morio ORS OLORE TOMO ce OR aaa ele OLOLO ORO RIEO
M. picicornis COMTOO NT a Ose ae OLOTOLO ORIG
M. picinus LPL OLOTO RL ORO ROLO RLS OF Seat is OL esr is tate
M. picipes OFZ OORT 1 e ee OM OLO FI tials ee OrO OO RO maa
. Body shape. [0] slender; [1] robust.
. Eyes. [0] not attaining superior edge of head; [1] attaining superior edge of head; [2] surpassing superior
edge of head. Treated as additive.
. Ocelli. [0] not placed on a tubercle; [1] placed on a tubercle.
. Lateral tubercle on neck. [0] present; [1] absent.
. Metallic shine in pronotum. [0] absent; [1] present.
. Pronotal granulations on anterior lobe. [0] absent; [1] present.
. Pronotal granulations on posterior lobe. [0] absent; [1] present.
. Pronotal sulci. [0] distinct; [1] not distinct.
. Pronotal lateral internal sulci. [0] distally united; [1] medially united; [2] not united. Treated as non-additive.
Lateral margin of pronotum. [0] carina absent; [1] carina present at entire length.
Scutellum coloration. [0] unicolorous; [1] bicolored.
Spongy fossa. [0] not occupying distal third of fore and hind tibiae; [1] occupying distal third of fore and
hind tibiae.
Female hemelytra. [0] macropterous; [1] brachypterous.
Hemelytra. [0] surpassing apex of abdomen; [1] not surpassing apex of abdomen.
Body color, [0] not uniform; [1] uniform.
Hemelytral pale stripe on corium and clavus. [0] present; [1] absent.
Color of fore femora. [0] bicolored; [1] unicolorous.
Color of hind femora. [0] unicolorous; [1] bicolored.
Color of fore tibiae. [0] unicolorous; [1] bicolored.
Color of mid tibiae. [0] unicolorous; [1] bicolored.
Connexivum. [0] dorsally visible; [1] dorsally not visible.
Connexivum color. [0] unicolorous; [1] bicolored.
Parameres shape. [0] subrectangular; [1] subtriangular.
Gonocoxite IX internal edge hairs. [0] thin; [1] thin and thick.
Gonocoxite IX unsclerotized area close to inner margin. [0] absent; [1] present.
Shape of IX and X tergites. [0] subquadrangular; [1] subrounded.
Intersegmental line of tergites IX and X. [0] not entire; [1] entire.
Intersegmental line. [0] not strongly sclerotized; [1] strongly sclerotized.
lin, Berlin, Germany; Zoological Museum,
University of Helsinki, Helsinki, Finland;
and the private collections of D. Carpintero,
Argentina; L. Jir6n, Costa Rica; and the late
J. Maldonado-Capriles, Puerto Rico.
Melanolestes constitutes a monophyletic
group that is distinguished from other Peira-
tinae by the spongy fossa occupying distal
third of fore and hind tibiae, body of uni-
formly dark color, ocelli placed on a tubercle,
pronotal granulations on anterior lobe, he-
melytra lacking a pale stripe on the corium
and clavus, hind femora unicolorous, and in-
tersegmental line of tergites [IX and X entire.
The nine species currently assigned to
the genus are considered as terminal taxa:
M. argentinus Berg, M. degener (Walker),
M. goiasensis Coscar6n and Carpintero, M.
lugens Coscar6n and Carpintero, M. minu-
tus Coscaron and Carpintero, M. morio (Er-
VOLUME 99, NUMBER 1
Peirates
Figs
= dotted rectangles.
ichson), M. picicornis (Stal), M. picinus
Stal, and M. picipes (Herrich-Schaeffer).
The data matrix and the 28 characters
used in this study are detailed in Table 1.
The data were analyzed with Hennig86 ver-
sion 1.5 (Farris 1988), applying the implicit
enumeration (ie*) option for calculating the
shortest trees. Consistency (CI) and reten-
tion (RI) indices were calculated excluding
uninformative characters (autapomorphies
and synapomorphies of the genus). We used
the successive weighting procedure in Hen-
nig86, that calculates weights from the best
fits of the characters on the most parsimo-
nious cladograms using rescaled consisten-
cies (products of the character consistency
and the character retention index). These
products are scaled in the range 0-10, and
the weighting procedure is repeated succes-
sively until the cladograms no longer
change (Farris 1989). CLADOS version 1.1
(Nixon 1992) was used for examination of
character distributions.
RESULTS AND DISCUSSION
The analysis using equal weights yielded
20 equally parsimonious cladograms, each
Cladogram of the species of Melanolestes.
M. picinus
M. degener
Synapomorphies = solid black rectangles; homoplasies
with 34 steps, CI = 0.52, and RI = 0.44.
When the successive weighting procedure
was applied, one minimum-length clado-
gram was selected from the original ones
after the second round of weighting, with
77 steps, CI = 0.84, and RI = 0.87 (Fig.
1). Values for the number of steps, consis-
tency index (ci), retention index (ri), and
weight (ri X ci X 100) in the weighted trees
are listed in Table 2. The phylogenetic se-
quence from the basal to the most distal
species is as follows: M. lugens, M. minu-
tus, M. picicornis plus M. picipes, M. mo-
rio, M. picinus plus M. goiasensis, and M.
argentinus plus M. degener.
Several conclusions can be deduced by
comparing the cladogram obtained with the
areas inhabited by the species of Melano-
lestes (Fig. 2):
(1) the more basal species (M. lugens) is
restricted to the Paranaense province of the
Amazonian domain;
(2) M. minutus is restricted to the Cha-
coan domain;
(3) the more widespread species M. mo-
rio and M. argentinus are among the most
distal species of the cladogram;
58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1000 2000 Km
Fig. 2. Geographical distribution of the species of Melanolestes, with the cladogram superimposed. a, M.
lugens; b, M. minutus; c, M. picicornis; d, M. picipes; e, M. morio, f, M. picinus; g. M. goiasensis; h, M.
argentinus; i, M. degener.
VOLUME 99, NUMBER 1
Table 2. Character consistencies and retention in-
dices are the best fits of the 20 most parsimonious
cladograms obtained applying ie. Final weights were
obtained after the second round of the successive
weighting procedure.
Number
of Consistency Retention Weight Final
Character Steps Index (ci) Index (ri) (ri X ci X 100) Weight
1 1 1.0 1.0 100 100
2 4 0.5 0.6 30 0
3 1 1.0 1.0 100 0)
4 i 1.0 1.0 100 100
5 2 0.5 0.5 2) 25
6 DD 0.5 0.5 DS 25
7 2 1.0 1.0 100 0
8 2 0.5 0) 0 10)
9 2 0.5 0) 0 0)
10 1 1.0 1.0 100 100
11 i 1.0 1.0 100 100
12 | 1.0 1.0 100 100
13 I 1.0 1.0 100 100
14 1 1.0 1.0 100 100
15 1 1.0 1.0 100 100
16 1 1.0 1.0 100 100
7 1 1.0 1.0 100 100
18 ] 1.0 1.0 100 100
19 1 1.0 1.0 100 100
20 1 1.0 1.0 100 100
21 1 1.0 1.0 100 100
22 1 1.0 1.0 100 100
28 2 0.5 10) 0) 0)
24 2 0.5 0.5 25 0)
25 1 1.0 1.0 100 100
26 1 1.0 1.0 100 33
2 1 1.0 1.0 100 100
28 2 0.5 0.5 25 DS
(4) the two Nearctic species (M. picicor-
nis and M. picipes) are sister-taxa.
These results corroborate, in part, our
previous study (Morrone and Coscar6n
59
1996), because the Chacoan species M.
minutus 1S one of the most basal species,
whereas the Amazonian species are among
the most distal species. Because M. pici-
cornis and M. picipes are sister-taxa, a sin-
gle dispersal event accounts for the pres-
ence of Melanolestes in the Nearctic.
ACKNOWLEDGMENTS
We thank Thomas J. Henry and one
anonymous reviewer for the critical reading
of the manuscript. Our study was supported
by grants from the University of Helsinki
to the senior author and grant 4662-91 of
the National Geographic Society to the ju-
nior author. Support from the Consejo Na-
cional de Investigaciones Cientificas y Téc-
nicas (CONICET), Argentina, to which the
authors belong, is gratefully acknowledged.
LITERATURE CITED
Cabrera, A. L. and A. Willink. 1973. Biogeografia de
América Latina. Monografia 13, Serie de Biolo-
gia, OEA, Washington D.C.
Coscaron, M. del C. and D. L. Carpintero. 1993. Re-
vision of the genus Melanolestes Stal (Heterop-
tera: Reduviidae, Peiratinae). Entomologica Scan-
dinavica 24(4): 361-381.
Farris, J. S. 1988. Hennig86 reference. Version 1.5.
Published by the author, New York.
—. 1989. The retention index and the rescaled con-
sistency index. Cladistics 5: 417—419.
Morrone, J. J. and M. del C. Coscar6n. 1996. Distri-
butional patterns of the American Peiratinae (Het-
eroptera: Reduviidae). Zoologische Mededelingen
Leiden.
Nixon, K. C. 1992. CLADOS ver. 1.1. IBM PC-com-
patible character analysis program. Published by
the author, Port Jefferson, New York.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 60-66
AULACIDAE (HYMENOPTERA) OF SRI LANKA
DAVID R. SMITH
Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart-
ment of Agriculture, % National Museum of Natural History, MRC-168, Washington, DC
20560, U.S.A.
Abstract.—Three species of Aulacidae from Sri Lanka are keyed, described, and illus-
trated: Pristaulacus flavipennis (Cameron), P. signatus (Shuckard), and P. krombeini, n.
sp. Lectotypes for Aulacus flavipennis Cameron and Aulacus magnificus Schletterer are
designated.
Key Words:
Three species of the family Aulacidae are
known from Sri Lanka, all of which were
among collections of the Smithsonian Cey-
lonese Insect Project of the 1970’s. Two are
species previously described from Sri Lan-
ka, and one was undescribed. All are in the
genus Pristaulacus Kieffer. About 20 spe-
cies of Aulacidae have been described from
the Oriental Region, and six of them are
from northern India, but the species from
Sri Lanka are distinct.
Species of Aulacidae are parasitoids of
wood-boring Coleoptera (especially Cer-
ambycidae and Buprestidae) and Hymenop-
tera (Xiphydriidae). Hosts are not known
for the Sri Lankan species, and there are
only two host records from southeastern
Asia. Pristaulacus beesoni Turner, de-
scribed from **Thano, Siwalik Hills, United
Provinces, India’’ was “‘taken emerging
from a Sal log” (Turner 1922), and a form
described as Pristaulacus nigripes var. du-
porti Kieffer from Viet Nam was ‘‘Obtenu
en abondance des larves de Xylotrechus
quadripes |Cerambycidae]”’ (Kieffer 1921).
KEY TO AULACIDAE OF SRI LANKA
1. Head, mesosoma, and legs mostly yellow;
wings yellowish, forewing sometimes with
Pe
Aulacidae, Pristaulacus, Sri Lanka, parasitoids
spot below stigma and apical margin black
(Fig. 4), or entirely blackish apical to stigma;
mesonotum reticulate (Fig. 2); head in dorsal
view elongate behind eyes (Fig. 1)
BS Can ee eae CAboe P. flavipennis (Cameron)
Head and body nearly all black, at most with
only antennal scape and pedicel and fore- and
midlegs reddish; forewing hyaline except for
black substigmatal spot and black spot at apex
or with anterior half black; mesonotum with
transverse ridges (Figs. 6, 10); head in dorsal
view short and narrowing behind eyes (Figs. 5,
Oy yea meet ete chad aye 2.4 ae ee
Forewing hyaline with infuscated spot below
stigma (Fig. 8); antennal scape and pedicel and
fore- and midlegs reddish; mesosoma with
rather dense silvery hairs obscuring sculpture
(Fig. 7); anterolateral lobes of mesonotum pro-
jecting forward in dorsal and lateral views
(Figs. 6, 7)
Forewing infuscated on anterior half, posterior
part nearly hyaline (Fig. 12); head, body, and
legs black, at most with mandible, tegula, and
first metasomal segment reddish to brownish;
mesosoma with relatively sparse hairs which
do not obscure sculpture (Fig. 11); anterolateral
lobes of mesonotum scarcely projecting for-
ward in dorsal view (Fig. 10), projecting up-
ward in lateral view (Fig. 11)
PAA Pee Tee P. krombeini, new species
Pristaulacus flavipennis (Cameron)
(Figs. 1—4)
Aulacus flavipennis Cameron 1887: 134
a eg dr P. signatus (Shuckard)
VOLUME 99, NUMBER 1
61
Figs. 1-4. Pristaulacus flavipennis. 1, Head, dorsal. 2, Mesosoma, dorsal. 3, Mesosoma, lateral. 4, Wings.
1-3, Specimen from Kegalla District. 4, Lectotype of P. magnificus.
(So): —Kietier 19253715372 (in key, ¢ fe-
described).
Pristaulacus flavipennis: Turner 1919:
386 (syn.: magnificus Schletterer).—Hedi-
exe 1959278:
Aulacus magnificus Schletterer 1890:
AO SSOO Pt 2201 SIC Ls "s:).
Pristaulacus magnificus: Kieffer 1900:
338.—Kieffer 1902: 13.—Kieffer 1912:
380, 388 (in key; d and @ redescribed).
Female.—Length excluding ovipositor,
16-17 mm; forewing length, 14 mm; ovi-
positor length, 16-17 mm. Antenna black.
Head yellow; apex of mandible black. Me-
sosoma yellow. Legs yellow with hindleg
black except base and inner surface of coxa
(lectotype of magnificus) or first segment of
trochanter, apical third of outer surface of
tibia, and tarsus black (specimen from Ke-
galla District). Metasoma, except base and
dorsal surface of first segment black (lec-
totype of magnificus) or orange with apical
third of metasomal terga 2—6 blackish
(specimen from Kegalla District); sheath
black. Wings yellow; forewing with stigma
black and large spot below stigma and ap-
icoventral margin black (lectotype of mag-
nificus; Fig. 4) or yellow with basal portion
of stigma blackish and very faint darker
spot below stigma (specimen from Kegalla
District). Head from above elongated be-
hind eyes (Fig. 1), shining and impunctate;
62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
occipital carina very short. Propleuron shin-
ing and impunctate. Front margin of pro-
notum with a forward projecting triangular
tooth just above its midlength; mesonotum
with reticulate sculpture; front margin of
mesonotum rounded, slightly lobate, not
overhanging pronotum in lateral view (me-
sosoma as in Figs. 2, 3). Hindcoxa with dis-
tinct, coarse cross wrinkles laterally and
posteriorly. Tarsal claws with 5 teeth (in-
cluding apical tooth) and small basal lobe.
Ovipositor length about 1.2 forewing
length.
Male.—Length, 15 mm. Color similar to
lectotype female (magnificus), except
smaller substigmatal spot of forewing,
which is about half width of stigma and
does not enter cubital cells (paralectotypes
of flavipennis and magnificus), or with apex
of forewing from base of stigma black (lec-
totype of flavipennis). Structure as for fe-
male.
Records.—SRI LANKA: Kandy District,
Dikoya, 06°52'N, 80°30’E (types of A. fla-
vipennis, spelled ““‘Dekaya’’ by Cameron,
1887). Kegalla District, Kitulgala, Makan-
de Mukalana, 3-4-11-1979, K.V. Krombein,
PB. Karunaratne, T. Wijesinhe, S. Siriwar-
dane, T. Gunawardane (1 2).
Types.—Aulacus flavipennis Cameron
was described from two males from “De-
kaya, Ceylon (Mr. George Lewis)’’ The
specimens are in The Natural History Mu-
seum, London. The colors of the two spec-
imens are slightly different, and Cameron
called them “‘two forms of this species.”
One specimen is labeled “B.M. TYPE
HYM 3.a.139’’, and this is hereby desig-
nated lectotype. The specimen lacks anten-
nae, except for the basal three segments of
the left antenna, hindlegs, and the metaso-
ma, except for the first segment. The other
specimen, the second “‘form”’ described by
Cameron, lacks a type label and is consid-
ered a paralectotype.
Schletterer described Aulacus magnificus
from a female and a male. He stated “‘Type
im kaiserl. naturhistorischen Hofmuseum
zu Wien (2) und im kGnigl. naturhistorisch-
en Museum zu Berlin (d).”” I have exam-
ined both specimens. The lectotype here
designated is the female at the Naturhisto-
risches Museum Wien, Austria, labeled
‘Felder [spelling ?], Ceylon, 1861,” ““mag-
nificus det. Schlett.”’, and a red label ““TY-
PUS.”’ The specimen is in fair condition
with the following parts missing: right an-
tenna beyond basal 4 segments, left antenna
beyond basal 6 segments, right forewing,
right foreleg, and left midleg. The male, a
paralectotype, is at the Zoologisches Mu-
seum an der Humboldt-Universitat zu Ber-
lin, Germany, labeled “‘Ceylon, Nietner.”
“11548,” “‘type’’ [red label], “‘magnificus
Schlett.,’’ and “‘Zool. Mus. Berlin.” It is in
good condition.
Discussion.—Even though there are col-
or differences in the wings, metasoma, and
hindlegs among the specimen from Kegalla
District (female), the two forms of A. fla-
vipennis Cameron (males), and the two
specimens of A. magnificus Schletterer (fe-
male and male), I regard these as probable
variation or sexual differences and treat
them as the same species. The structure, es-
pecially sculpture, of all is similar. Much
more study material will be necessary to
determine if this is a species complex with
perhaps more than one species involved.
The specimen collected in Kegalla District
is the palest: yellow with antenna, apex of
mandible, apical portion of terga 2—6,
sheath, ovipositor, first segment of hindtro-
chanter, apical third of outer surface of
hindtibia, and all hindtarsus black, and the
forewing is yellow with only a faint black
area beneath the stigma. The lectotype of
A. flavipennis has most of the metasoma
and hindtibia black and the forewing with
the apex beyond the basal end of the stigma
mostly black. The paralectotype of A. fla-
vipennis is similar to the lectotype except
the forewing has a large spot below the
stigma and apical margin black, and the lec-
totype female and paralectotype male of A.
magnificus Schletterer are similar to the
paralectotype of A. flavipennis.
VOLUME 99, NUMBER I
Figs. 5-8.
Pristaulacus signatus (Shuckard)
(Figs. 5-8)
Aulacus signatus Shuckard 1841: 124.—
Westwood 1844: 268.—Schletterer 1890:
509 (= ? stigmaticus Westwood).—Kieffer
1902: 12 (? Aulacus).—Kieffer 1912: 373
(repeats Shuckard’s description).—Hedicke
193919:
Pristaulacus signatus: Kieffer 1904: 455
(? Pristaulacus).
Female.—Length, excluding ovipositor,
9-11 mm, forewing length, 8-10 mm; ovi-
positor length, 8-10 mm. Black except for
following: mandible dark orange with apex
reddish brown; anterior half or less of clyp-
eus sometime brownish; antennal scape or
Pristaulacus signatus. 5, Head, dorsal. 6,
Mesosoma, dorsal. 7, Mesosoma, lateral. 8, Wings.
scape and pedicel and fore- and midlegs be-
yond trochanters dark reddish; hindleg be-
yond coxae sometimes brownish; tegula
brownish to reddish brown; first gastric seg-
ment sometimes partly pale reddish to or-
ange. Wings hyaline; forewing with small
infuscated spot below stigma (Fig. 8). Head
from above short and narrowing behind
eyes (Fig. 5); shining with widely scattered
punctures and silvery hairs, these both more
dense between ocelli and antennae; occipi-
tal carina short. Propleuron shining with
scattered punctures, similar to top of head.
Pronotum with a forward projecting trian-
gular tooth just above its midlength; antero-
lateral angles of mesonotum protruding an-
teriorly in dorsal and lateral views and
64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
mesonotum with coarse transverse ridges;
dense silvery hairs somewhat obscuring
sculpture, especially laterally (mesosoma as
in Figs. 6, 7). Hindcoxa with distinct,
coarse transverse wrinkles laterally and
posteriorly. Tarsal claws with 4 teeth, in-
cluding apical tooth, and small basal lobe.
Ovipositor length about as long as forewing
length.
Male.—Length, 8-10 mm. Color and
structure as for female.
Records.—SRI LANKA: Amparai Dis-
trict, Ekgal Aru, 12-VI-1976, K.V. Krom-
bein, P.B. Karunaratne, S. Karunaratne (1
2); Ekgal Aru, 100 mtrs, Reservoir Jungle,
19-22-11-1977, K.V. Krombein, P.B. Karu-
naratne, P. Fernando, D.W. Balasooryia (1
2), 11-12-VI-1976, K.V.Krombein, PB.
Karunaratne, S. Karunaratne (1 2); Lahu-
gala Tank, 14-15-VI-1976, K.V.Krombein,
PB. Karunaratne, S. Karunaratne (1 °). An-
uradhapura District, Padaviya Tank, 180
ft., 20-21-V-1976, K.V.Krombein, P.B. Ka-
runaratne, S. Karunaratne, D.W. Balasoor-
yia (1 2, 1 d); Padaviya, 180 ft., Irrigation
Bungalow, 18-V-1976, K.V. Krombein, P.B.
Karunaratne, S. Karunaratne, D.W. Bala-
sooryia (1 2). Colombo District, Labuga-
ma, 23-24-VI-1975, S.L. Wood & J.L. Petty
(2 2). Kandy District, Hasalaka, 140-170
m, 15-17-IX-1977, K.V. Krombein, P.B.
Karunaratne, T. Wijesinhe, M. Jayaweera (1
2). Kegalla District, Kitulgala, Makande
Mukalana, 3-4-II-1979, K.V. Krombein,
P.B. Karunaratne, T. Wijesinhe, S. Siriwar-
dane, T. Gunawardana (1 d). Polonnaruwa
District, 25 mi NW Polonnaruwa, 11-VI-
1975; Sil» Woodrc& Tk PettyaC +2) aN:
Central Province, Ritigala Nat. Reserve, 8
mi NW Habarana 8-II-1962. Loc. 546:],
swept on grass in forest, Lund University
Ceylon Expedition 1962, Brinck-Anders-
son-Cederholm (1 2). Puttalam District, 17
mi SE Puttalam, 18-VI-1975, S.L. Wood &
J.L Petty (2 2). Ratnapura District, Uggal-
kaltota, 350’, 20-VI-1976, K.V. Krombein,
PB. Karunaratne, S. Karunaratne (1 Q);
Gilimale, Induruwa Jungle, 5-7-11-1977,
K.V. Krombein, P. Fernando, D.W. Bala-
sooryia, V. Gunawardane (1 2). Trincom-
alee District, China Bay Ridge Bungalow,
0-50 feet, 24-25-VIH-1978, K.V. Krombein,
T. Wijesinhe, V. Kulasekare, L. Jayawick-
rema (1 2, 2 d); Trincomalee, China Bay
Ridge Bungalow, 25-50 ft., 26-II-1979,
K.V. Krombein, T. Wijesinhe, S. Siriwar-
dane, L. Jayawickrema, T. Gunawardane (2
2), 0-100’, 13-17-V-1976, K.V. Krombein,
PB. Karunaratne, S. Karunaratne, D.W.
Balasooryia (1 2), 16-17-V-1976, same
collectors (1 2); Trincomalee, China Bay,
1-30 m, 8-11-X-1977, collected near ridge
bungalow, K.V. Krombein, P.B. Karunarat-
ne, P. Fernando, T. Wijesinhe, M. Jayaweera
(1 ¢), in Malaise trap, same date and col-
lectors (1 @).
Type.—The type is presumably lost. It
was not found in The Natural History Mu-
seum, London (A. Shinohara, correspon-
dence).
Discussion.—Shuckard’s type was not
found. His description is very brief: “*Ni-
ger: scapo antennarum pedibusque 4 anticis
rufo-testaceis: alis hyalinis, macula ad stig-
ma brunnea. Long. 5% lin. Exp. alar. 9%
lin.’ The sex is not given and cannot be
determined since both sexes are colored
similarly. The specimens from Sri Lanka
agree with Shuckard’s description, and I re-
gard them as A. signatus. This species ap-
pears to be the most widely distributed spe-
cies in Sri Lanka and is easily separated
from the other two species by the black col-
or with the reddish scape, pedicel, and fore-
and midlegs and structural characters as
given in the key and as illustrated.
Pristaulacus krombeini Smith,
new species
(Figs. 9-12)
Female.—Length, excluding ovipositor,
16 mm; forewing length, 13 mm; ovipositor
length, 15 mm. Black; tegula reddish
brown; anterior margin of clypeus, mandi-
ble except apex, tarsi, and lateral and ven-
tral portions of first metasomal segment
slightly brownish. Forewing with anterior
VOLUME 99, NUMBER 1
oO
Nn
Figs. 9-12.
half black, posterior half of forewing and
hindwing more hyaline to slightly blackish;
veins and stigma black (Fig. 12). Head from
above short and narrowing behind eyes
(Fig. 9); with widely scattered fine punc-
tures and silvery hairs, each more dense be-
tween ocelli and antennae; occipital carina
short. Propleuron shining with scattered
punctures, similar to top of head. Pronotum
with a forward projecting triangular tooth
just above its midlength; anterolateral an-
gles of mesonotum scarcely protruding and
rounded in dorsal view, directed upward in
lateral view, mesonotum with coarse trans-
verse ridges; silvery hairs on mesosoma not
obscuring sculpture (mesosoma as in Figs.
10, 11). Hindcoxa with coarse transverse
Pristaulacus krombeini. 9, Head, dorsal. 10, Mesosoma, dorsal. 11, Mesosoma, lateral. 12, Wings.
wrinkles laterally, posterior surface almost
devoid of wrinkles. Tarsal claws with 4
teeth, including apical tooth, and a small
acute basal lobe. Ovipositor length about
1.2 X forewing length.
Male.—Unknown.
Holotype.—Female, labeled “Sri Lanka:
Tri. [Trincomalee] Dist., Tennamaravadi,
18-V-1976, K.V. Krombein, P.B. Karunar-
atne, S. Karunaratne, D.W. Balasooryia.”’ In
the National Museum of Natural History,
Washington, D.C.
Etymology.—Named for my colleague,
Dr. Karl V. Krombein, Department of En-
tomology, Smithsonian Institution, Wash-
ington, D.C.
Discussion.—The forewing with the an-
66 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
terior half infuscated and the posterior half
lighter black to hyaline, entirely black col-
oration, and structure of the mesonotum as
shown in Figs. 10 and 11 separate this spe-
cies from other Aulacidae described from
southeastern Asia. A review of the descrip-
tions of other species described from south-
eastern Asia, especially those from northern
India and Indochina, revealed various dis-
tinguishing features from P. krombeini, es-
pecially combinations of coloration, wing
maculation, and mesonotum structure.
These taxa include P. beesoni Turner, P. ir-
idipennis (Cameron), P. nigripes Kieffer, P.
rufobalteatus Cameron, P. rufobalteatus
leviceps Kieffer, and P. bituberculatus
(Cameron) from the Himalayas of northern
India, and P. emarginaticeps Turner, P. ex-
cisus Turner, P. nigripes var. duporti Kief-
fer, P. tuburculiceps Turner, and P. tonki-
nensis Turner from Indochina.
ACKNOWLEDGMENTS
I thank K.V. Krombein, Smithsonian I[n-
stitution, for allowing study of material
from the Ceylonese Insect Project, and the
following for loaning types and other ma-
terial: R. Danielsson, Museum of Zoology,
Lund University, Lund, Sweden; S. Schédl,
Naturhistorisches Museum Wien, Austria;
and FE Koch, Zoologisches Museum an der
Humboldt-Universitét zu Berlin, Germany.
I am especially grateful to A. Shinohara,
National Science Museum, Tokyo, Japan,
who kindly compared photos with types
and provided notes on the types of A. fla-
vipennis during his visit to The Natural His-
tory Museum, London, in 1995.
LITERATURE CITED
Cameron, P. 1887. Descriptions of one new genus and
some new species of parasitic Hymenoptera. Pro-
ceedings of the Manchester Literary and Philo-
sophical Society 26: 117—136.
Hedicke, H. 1939. Aulacidae. /n Hedicke, H., ed., Hy-
menopterorum Catalogus, Pars 10. Dr. W. Junk,
Gravenhage. 28 pp.
Kieffer, J.-J. 1900. Note sur le genre Pristaulacus
Kieff. [Hymén.]. Bulletin de la Société Entomo-
logique de France 1900: 338-339.
1902. Hymenoptera, Fam. Evaniidae. In
Wytsman, P., Genera Insectorum, Fascicule 2, 13
pp., | plate. Bruxelles.
1904. Les Evaniides. Jn André, E., Species
des Hyménopteéres d’ Europe et d’ Algérie, Volume
7, part 2, pp. 347—482. Paris.
1912. Hymenoptera, Ichneumonidae, Evani-
idae. Das Tierreich, Berlin, Volume 30, I-XIX +
431 pp.
. 1921. Sur divers Hyménopteres destructeurs
des Cérambycides nuisibles au Caféier et au Bam-
bou. Bulletin Agricole de I’ Institut Scientifique de
Saigon 3: 129-140.
Schletterer, A. 1890. Die Hymenopteren-Gruppe der
Evaniiden. Annalen des K. K. Naturhistorischen
Hofmuseums (Separatabdruck aus Band IV), pp.
373-546, plates XIX-XXII.
Shuckard, W. E. 1841. Art. XXII.—On the Aulacidae,
a family of Hymenoptera pupivora; and that 77i-
gonalys is one of its components: with the de-
scription of a British species of this genus, and
incidental remarks upon their collateral affinities.
Entomologist 1: 115-125.
Turner, R. E. 1919. On Indo-Chinese Hymenoptera
collected by R. Vitalis de Salvaza.—lII. Annals
and Magazine of Natural History (9) 4: 385-395.
. 1922. XXXIV.—New Evaniidae and Bracon-
idae in the British Museum. Annals and Magazine
of Natural History (9) 10: 270-281.
Westwood, J. O. 1844 (1843). XXXVIII. On Evania
and some allied genera of Hymenopterous insects.
Transactions of the Entomological Society of Lon-
don 3: 237-278, plates XITV—XV.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 67-88
NOTES ON LIFE HISTORIES AND DESCRIPTIONS OF ADULTS AND
IMMATURE STAGES OF PROCECIDOCHARES KRISTINEAE AND P. LISAE
NEW SPECIES (DIPTERA: TEPHRITIDAE) ON AMBROSIA SPP. IN
SOUTHERN CALIFORNIA
RICHARD D. GOEDEN AND JEFFREY A. TEERINK
Department of Entomology, University of CA, Riverside, CA 92521, U.S.A.
Abstract.—Procecidochares kristineae Goeden, n. sp., and P. lisae Goeden, n. sp.,
are described and figured as adults and distinguished from each other by their wing
patterns, chaetotaxy, and genal color and by these and additional characters from all other
North American congeners. Sexually dimorphic wing patterns in P. lisae are reported for
the first time from this genus. First through third-instar larvae and puparia also are de-
scribed and figured for both species, but few morphological differences were found. Pro-
cecidochares kristineae and P. lisae are strictly monophagous tephritid flies forming ax-
illary bud galls on branches of the native, shrubby, xerophytic ragweeds, Ambrosia du-
mosa (Gray) Payne and A. eriocentra (Gray) Payne (Asteraceae), respectively. Both are
circumnatal species that overwinter as first instars within incipient galls in southern Cal-
ifornia. Further gall and larval development and host-plant regrowth and reproduction are
stimulated concurrently by winter rainfall. Reproduction by P. kristineae is rarely biennial,
usually univoltine or bivoltine, and rarely trivoltine, depending on local yearly rainfall
patterns; whereas, P. /isae usually is univoltine or bivoltine.
Key Words: Insecta, Procecidochares, Ambrosia, ragweed, biology, galls, taxonomy of
adults and immature stages, sexual dimorphism, parasitoid
Two undescribed species of Procecido-
chares (Diptera: Tephritidae) were detected
during faunistic surveys of native rag-
weeds, Ambrosia spp. (Asteraceae: Ambro-
Siinae), in southern California by Goeden
and Ricker (1976a, b). The life history of
one species forming galls on A. dumosa
(Gray) Payne was studied by Silverman and
Goeden (1980), but it has remained un-
named until now. In this paper, both te-
phritid species are named, their adult and
immature stages are described and illustrat-
ed, and new life-history information on
each is reported.
MATERIALS AND METHODS
Our field studies on the tephritid infest-
ing A. dumosa were conducted near sea lev-
el in the low-elevation Colorado Desert (=
northern Sonoran Desert) in southern Cali-
fornia (Munz 1974) at various locations in
Imperial, Riverside, and San Diego counties
listed by Silverman and Goeden (1980) or
mapped by Goeden and Ricker (1976a).
Our principal study site for the fly on A.
eriocentra (Gray) Payne was Mountain
Pass at 1430-m elevation, San Bernardino
Co., in the high-elevation, Mojave Desert;
although this tephritid also was reared from
galls obtained from surrounding locations
named below in northeastern San Bernar-
dino Co. during 1970—71 faunistic surveys
(Goeden and Ricker 1976b). Galls contain-
ing eggs, larvae and puparia were sampled
most recently from A. dumosa near Snow
68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Creek at 370-m elevation in the dry San
Gorgonio River bed, Riverside Co., during
February—April, 1993 and 1994, and from
A. eriocentra at Mountain Pass during Feb-
ruary—May, 1991-1995. Excised galls were
transported in cold-chests in an air-condi-
tioned vehicle to the laboratory and stored
under refrigeration for subsequent dissec-
tion, photography, description, and mea-
surement. Twenty-one and 19 larvae and 4
and 3 puparia dissected from galls on A.
dumosa and A. eriocentra, respectively,
were preserved in 70% EtOH for scanning
electron microscopy (SEM). All other fully
grown larvae and puparia from A. eriocen-
tra were placed in separate, glass shell vials
stoppered with absorbant cotton and held in
humidity chambers at room temperature for
adult and parasitoid emergence. Specimens
for SEM later were hydrated to distilled wa-
ter in a decreasing series of acidulated
EtOH. They were osmicated for 24 h, de-
hydrated through an increasing series of
acidulated EtOH, critically point dried,
mounted on stubs, sputter-coated with a
gold-palladium alloy, and studied with a
JEOL JSM C-35 SEM in the Department of
Nematology, University of California, Riv-
erside.
Most adults reared from isolated puparia
obtained from galls on A. eriocentra were
individually caged in 850-ml, clear-plastic,
screened-top cages with a cotton wick and
basal water reservoir and provisioned with
a strip of paper toweling impregnated with
yeast hydrolyzate and sucrose. These cag-
ings were used for longevity studies and
Oviposition tests in the insectary of the De-
partment of Entomology, University of Cal-
ifornia, Riverside, at 25 + 1°C, and 14/10
(L/D) photoperiod. Virgin male and female
flies obtained from emergence vials as well
as field-swept adults were paired in clear-
plastic petri dishes provisioned with a flat-
tened, water-moistened pad of absorbant
cotton spotted with honey (Headrick and
Goeden 1991) for direct observations, vi-
deorecording, and still-photography of their
courtship and copulation behavior.
Plant names used in this paper follow
Munz (1974); tephritid nomenclature and
adult terminology follow Foote et al.
(1993). Format used to describe the adults
follows Blanc and Foote (1961). Morpho-
logical terminology and telegraphic format
used to describe the immature stages follow
Goeden and Headrick (1992), Headrick and
Goeden (1990, 1993), Goeden et al. (1994a,
b, 1995a, b), Goeden and Teerink (1996a,
b, c), Headrick et al. (1995), and our other
works cited therein. Means + SE are used
throughout this paper. The holotypes, allo-
types, and five individually reared para-
types of each sex of both new species have
been deposited in the National Museum of
Natural History, Smithsonian Institution,
Washington, D.C. (USNM). Six additional,
individually reared paratypes of each sex of
both species also were deposited in the col-
lection of the California Academy of Sci-
ences (CAS). The holotype, allotype, and
22 paratypes used for measurements to de-
scribe P. kristineae originated from one col-
lection in 1980 at the Palm Springs study
site used by Silverman and Goeden (1980);
the same numbers of types were used to
describe P. lisae from one collection in
1981 at Mountain Pass. All remaining para-
types and voucher specimens not designat-
ed as paratypes and reared parasitoids of
both tephritids reside in the research collec-
tions of RDG; preserved specimens of lar-
vae and puparia are stored in a separate col-
lection of immature Tephritidae maintained
by JAT.
RESULTS AND DISCUSSION
TAXONOMY
The new species from A. dumosa de-
scribed below as P. kristineae is most sim-
ilar to P. stonei Blanc and Foote, which it
was first misidentified as (Silverman and
Goeden 1980). However, adults of these
two species are distinguished below mainly
on the basis of their wing characters, in-
stead of those often lacking in swept or pre-
served specimens, e.g., distribution patterns
VOLUME 99, NUMBER 1
of setae on the scutum, consistent with the
philosophy embodied in the key to U.S. and
Canadian species of Procecidochares in
Foote et al. (1993). The new species from
A. eriocentra described as P. lisae also is
distinguished below primarily by wing
characters. Accordingly, the following key
couplets replacing couplets 1 and 5 in the
key of Foote et al. (1993) enable one to
distinguish these two new species:
1. Pterostigma lighter brown in basal fifth to half
than in apical part; sometimes a second orbi-
talbrisilo mee eer eat cents. cheer ese la
— Pterostigma evenly brown; | pair orbital bris-
WES A ards ot 884 youn doen cede taker Ok eran 2.
la. Two orbital bristles, the second pair very
small and delicate; basal and discal bands
SCPalale wt ems ies alsa flavipes Aldrich
— One pair of orbital bristles; basal and discal
bands usually connected
1b. Gena with prominent dark brown spot at ven-
tral margin of eye; discal band not extended
posterior to vein A, + CuA, in male, usually
crossing it in female, but fading towards pos-
terior wing margin; basal and discal bands
broadly connected in cells c, sc, br, and bm
+ Bug abt OfdLc sei aA OTE ENS ee lisae Goeden, n. sp.
— Gena with ocherous to pale brown spot at
ventral margin of eye; discal band extending
distinctly to posterior wing margin in both
sexes; basal and discal bands narrowly con-
nected in cell sc or br . . kristinae Goeden, n. sp.
Procecidochares kristineae Goeden,
new species
(Figs. 1, 3-7)
Procecidochares stonei Blanc and Foote:
Wasbauer 1972: 7 (in part, Ambrosia du-
mosa host record).
Procecidochares n. sp.: Goeden and
Ricker 1976a: 49 (host record).—Silverman
and Goeden 1980: 283-288 (host, gall de-
scription, California distribution, biology,
behavior, seasonal history, parasitoids,
predators, gall inquiline).—Foote et al.
1993: 318 (taxonomic status)
Female (Holotype).—Head: In profile
0.6 to 0.7 times as long as high, face and
frons meeting at an angle of about 120°;
parafacial 0.75 times as wide as third an-
tennal segment; gena about 0.2 times as
high as eye, which is 0.5 to 0.6 as wide as
69
high; frons ocherous brown to yellow or
white, at vertex 1.4 to 1.6 times as wide as
eye in lateral view, 1.2 to 1.3 times as wide
as length from vertex to lunule; lunule half
as high as its width between the antennae;
face yellow to white, pollinose, concave,
but raised medially and protruding slightly
at middle of oral margin; antenna yellow,
pollinose, third segment sometimes ocher-
ous brown along anterior margin and apex,
arista ocherous brown to black, lightest ba-
sally. Usually 3-4 frontal bristles (rarely 5
or 6), all black; one pair of black orbital
bristles; black genal bristle slender, situated
immediately below lower curvature of eye;
all postoculars white.
Thorax: Pleuron mostly shining dark
brown to black, especially the katepister-
num, but with a very sparse pollinosity on
anepisternum; katepisternal bristle black,
most other pleural setae rather long and
white; wing base and anepimeron densely
silver pollinose over a dark brown to black
ground-color; lateral third of mesonotum,
including postpronotal lobe, shining dark
brown to black; a wide, silvery pollinose
stripe on a shiny black ground-color occu-
pying median third of mesonotum from an-
terior margin nearly to scutoscutellar suture,
slightly widened at transverse suture and in-
vested with short, white, blunt setae (some-
times appearing pale yellow) as follows:
scattered + uniformly over median polli-
nose strip, except for anterior half of pre-
sutural part of scutum, where confined to
center and margins of strip, also encircling
the lateral third of presutural part of scutum
including along the transverse suture, and
trailing posteriolaterad and posteriomediad
in separate rows 3 to 4 setae-wide, the outer
row narrowing and crossing the scutoscu-
tellar suture to end in a cluster of 5 to 12
setae at base of anterior scutellar bristle.
One dorsocentral bristle situated about half-
way between transverse suture and level of
the postsutural supra-alar bristle, and locat-
ed on margin of median pollinose area;
cluster of 5 to 8 short, white setae anterior-
ad of base of postsutural supra-alar bristle.
70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Right wings of Procecidochares kristineae:
Fig. 1.
(A) female; (B) male.
Scutellum shining dark brown to black, bul-
bous, two pairs scutellar bristles; postscu-
tellum dark brown to black, pollinose. Hal-
tere shaft ocherous yellow, slightly darker
than knob. Femora of fore, mid, and hind
legs dark brown but with yellow extremi-
ties, the remainder of legs yellow to ocher-
ous yellow. Wing pattern as in Fig. 1A, B;
pterostigma about 2.0 times as long as
wide, lighter in basal fifth to third; basal
and discal bands narrowly joined in cell br
and/or sc, subapical and apical bands usu-
ally joined in cell(s) r, and/or r,,, (Fig. 1A)
or sometimes narrowly separated (Fig 1B)
and only joined by darkened portions of
veins R,,, and R,,;; distance between
crossveins r-m and dm-cu measured along
vein M about equal to length of r-m.
Abdomen: Dark brown to black, each ter-
gum covered with white, flattened setae
progressively increasing in length posteri-
orly. Oviscape shining dark brown to black,
basal half slightly swollen, about half as
long as all terga, except the first, taken to-
gether.
Male (allotype)——Head: Like &, but
parafacial 0.67 times as wide as third an-
tennal segment; frons at vertex 1.1 to 1.3
times as wide as one eye, 1.1 to 1.4 times
as wide as distance from vertex to lunule;
lunule 0.5 to 0.7 times as high as its width
at antennal base. Thorax: Like 2°. Abdo-
men: Like 2, only external genitalia dark
brown to black.
Variation.—Examination of P. kristineae
specimens with mostly intact setation con-
firmed variation in numbers of frontal setae,
noted for the genus to vary from two to five
by Foote et al. (1993). The holotype and 19
? paratypes (68%) of a total of 28 2 types
examined had three pairs of frontal setae;
one pair of these setae was white in two &,
and one pair was reduced in length and
thickness in another 2. One @ had four
pairs of frontal setae; another 2 had five
pairs. The frontal setae were not strictly
paired in the remaining six @, as four @
had three and four frontal setae on different
sides of their heads; and each of two &,
four and five frontal setae located asym-
metrically. The allotype and 32 d paratypes
(64%) of a total of 50 d types examined
had three pairs of frontal setae. Three d
(6%) had four pairs of frontal setae. Among
the remaining 15 6d, nine (18%) had three
and four frontal setae on different sides of
their heads, four d (16%) each had three
and five frontal setae, and one d each had
four and five or four and six frontal setae
located asymmetrically.
Regarding the wings, the subapical and
apical bands also showed variation. The ho-
lotype and 14 @ paratypes (48%) of a total
of 31 2 types with intact wings had the
subapical and apical bands broadly joined
(Fig. 1A); 7 2 paratypes (23%) had the
subapical and apical bands narrowly joined
in at least one wing (in the other wing in 5
of these 2, these bands were narrowly sep-
arated, Fig. 1B); and in the remaining 9 2
(29%), these bands were narrowly separat-
ed in both wings. The allotype and 25 d
VOLUME 99, NUMBER 1
types (49%) of a total of 51 d paratypes
with intact, fully formed wings had the sub-
apical and apical bands broadly joined (Fig.
1A); 8 3d paratypes (16%) had these bands
narrowly joined in at least one wing (in the
other wing in 1 d, these bands were nar-
rowly separated); and in the remaining 18
3d paratypes (35%), these bands were nar-
rowly separated in both wings.
Diagnosis.—The two main morphologi-
cal characters distinguishing the adults of P.
kristineae are the pterostigma being lighter
brown in the basal fifth to third than in the
apical part (Fig. 1), in combination with one
pair of orbital setae. All types of P. kristi-
neae possessed these two characters. The
former character is shared only with P. fla-
vipes and P. lisae (see below); the latter
character distinguishes P. flavipes which
has two pairs of orbital setae (Foote et al.
1993). Procecidochares kristineae and P.
lisae are distinguished below. In the partial
key provided above, if the pterostigmal
character is missed P. kristineae will run to
P. stonei, the only other species with basal
and discal bands connected. Most P. kris-
tineae differ from P. stonei by having the
subapical and apical bands broadly to nar-
rowly joined.
Of 179 reared voucher specimens of P.
stonei with fully formed wings in the re-
search collection of RDG (Green et al.
1993), 14 (8%) had a pterostigma in at least
one wing that was light basally, but usually
this basal area was small and only partially
extended posteriorad across the cell, nor
were the subapical and apical bands in the
wings of these 14 flies joined. Similarly, an-
other 12 voucher specimens of P. stonei
had at least one wing with the subapical and
apical bands joined, but all of these flies
had pterostigmas that were evenly brown.
Additional biological and ecological char-
acteristics that distinguish P. kristineae and
P. stonei are discussed below.
Types.—Holotype, 2; 8 km NE of Palm
Springs, 250-m elevation, Riverside Co.,
Galiformiay 2641198 1:,.R..;D...Goeden,
coll.(hereafter, RDG, coll.); reared from ax-
val
illary bud gall on A. dumosa [at study site
of Silverman and Goeden (1980)] (USNM);
allotype, ¢6, same data as holotype
(USNM). Paratypes: CALIFORNIA: 12 ¢
and 24 2; same data as holotype (5 d and
5 2 to USNM, 6 ¢ and 6 2 to CAS). 4 6
and 3 2; same location data as holotype;
10.11.1971; RDG and D.W. Ricker, coll.
(hereafter, RDG & DWR, coll.). 1 2; Twen-
tynine Palms, San Bernardino Co.;
25.iii.1970; RDG & DWR, coll. 4 6 and 1
2; Llano, San Bernardino Co.; 30.iv.1970;
RDG & DWR, coll. 6 d and 1 2; Yucca
Valley, San Bernardino Co.; 4.111.1971;
RDG & DWR, coll. | 2; Borrego Springs,
San Diego Co., 1.11.1973; RDG & DWR,
coll. 5 ¢ and 7 2; Valliceto Valley, S end
of Smugglers Canyon at 442 m, San Diego
Co.; 11.11.1993; RDG and J.A. Teerink,
coll. (hereafter, RDG & JAT, coll.).5 ¢ and
4 2; Snow Creek at 370 m, Riverside Co.;
7.iv.1994; RDG & JAT, coll. 3 6 and 3 &;
Ocotillo, Imperial Co., 2.11.1995; RDG &
JAT, coll. (Remaining 39 d and 20 & para-
types along with numerous swept and/or
damaged-reared, voucher and nonvoucher
specimens examined and identified are held
in research collection of RDG).
Etymology.—Procecidochares kristineae
is named for my younger daughter, Kristine
Louise Gilbert (nee Goeden), mother of my
two grandsons, Samuel Vanderpoel Gilbert
V and Nikolaus Richard Gilbert.
Procecidochares lisae Goeden,
new species
(Figs. 2, 8-12)
Procecidochares n. sp.: Goeden and Ricker
(1976b): 927 (host record).
Female (holotype).—Head: In profile 0.5
to 0.6 times as long as high, face and frons
meeting at an angle of about 120°; parafa-
cial as wide as third antennal segment; gena
with prominent, shiny, dark-brown spot
from lower margin of eye to genal groove,
gena 0.1 to 0.2 times as high as eye, which
is 0.4 to 0.5 as wide as high; frons ocherous
yellow to white, at vertex 1.3 to 1.7 times
M2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
as wide as eye in lateral view, 1.3 to 1.6
times as wide as length from vertex to lu-
nule; lunule 0.4 to 0.6 times as high as its
width between the antennae; face pale yel-
low to white, pollinose, concave, but raised
medially; antenna mostly brown, pollinose,
apex of second segment and sometimes
posterior margin of third segment yellow,
arista ocherous yellow to brown, lightest
basally. Usually, 3—4 frontal bristles (rarely
2 or 5), all black; one pair of black orbital
bristles; genal bristle black, arising behind
brown part of genal margin, which is ex-
tension of brown spot (noted above) below
genal groove; all postoculars white.
Thorax: Pleuron mostly shining dark
brown to black, especially the katepister-
num, but with a very sparse pollinosity on
anepisternum; katepisternal bristle black,
most other pleural setae rather long and
white; wing base and anepimeron densely
silver pollinose over a dark brown to black
ground-color; lateral third of mesonotum,
including postpronotal lobe, shining dark
brown to black; a wide, silvery pollinose
stripe on a shiny black ground-color occu-
pying median third of mesonotum from an-
terior margin nearly to scutoscutellar suture,
slightly widened at transverse suture and in-
vested with short, white, blunt setae (some-
times appearing pale yellow) as follows:
scattered + uniformally over median polli-
nose strip, except for anterior half of pre-
sutural part of scutum, where confined to
center and margins of strip, also encircling
the lateral third of the presutural part of scu-
tum including along the transverse suture,
encircling the anterolateral two-thirds of the
scutum, and trailing posteriolaterad and
crossing the scutoscutellar suture to end in
a cluster of 8 to 18 setae at base of anterior
scutellar bristle. One dorsocentral bristle
situated just anterior to a line between the
postsutural supra-alars and slightly laterad
of the margin of median pollinose area.
Scutellum shining dark brown to black, bul-
bous, two pairs scutellar bristles; postscu-
tellum dark brown to black, pollinose. Hal-
tere shaft ocherous yellow, slightly darker
B
Right wings of Procecidochares lisae: (A)
Fig. 2.
female; (B) male.
than knob. Femora of fore, mid, and hind
legs dark brown but with ocherous yellow
extremities, the remainder of legs yellow to
ocherous yellow. Wing pattern as in Fig. 2A
pterostigma about 2.0 times as long as
wide, lighter in basal fifth to half; basal and
discal bands broadly joined in cells c, sc,
and br; discal band usually shortened,
crossing cell cua, along and distal to vein
CuA,, and nearly or just touching vein A,
+ CuA, growing faint if extending into anal
lobe; subapical and apical bands usually
separated, or sometimes narrowly joined in
cell(s) 1, and/or 1,5; distance’ between
crossveins r-m and dm-cu measured along
vein M about equal to length of r-m.
Abdomen: Dark brown to black, each ter-
gum covered with white, flattened setae
progressively increasing in length poste-
riorad. Oviscape shining dark brown to
black, basal half slightly swollen, about half
as long as all terga, except the first, taken
together.
VOLUME 99, NUMBER I
Male (allotype)—Head: Like @, but
about 0.6 times as long as high, face and
frons meeting at an angle of about 110°;
parafacial 0.67 times as wide as third an-
tennal segment; eye 0.5 to 0.6 times as wide
as high; frons at vertex 1.1 to 1.5 times as
wide as eye, 1.3 to 1.7 times as wide as
length from vertex to lunule; lunule 0.5 to
0.7 times as high as its width at antennal
base. Thorax: Like 2, but discal band of
wing rarely extending into anal lobe and
usually ending before and not touching vein
A, + CuA,; subapical and apical bands usu-
ally separated, but sometimes narrowly or
widely joined in cell(s) r, and/or r,,, (Fig.
2B). Abdomen: Like @, only external gen-
italia dark brown to black.
Variation.—Examination of P. lisae
specimens with mostly intact setation fur-
ther confirmed variation in numbers of
frontal setae for the genus (Foote et al.
1993). The holotype and 30 @ paratypes
(50%) of a total of 62 2 types examined
had three pairs of frontal setae. Six @ para-
types (13%) each had four pairs of frontal
setae; another @ had five pairs. The frontal
bristles were not paired in the remaining 22
2, as 19 2 had three and four frontal setae
on different sides of their heads; two 9 had
two and three frontal setae located asym-
metrically; and one @ had three and six
frontal setae so located. The allotype and
25 6d paratypes (53%) of a total of 47 ¢
types examined had three pairs of frontal
setae. Eight d paratypes (13%) each had
four pairs of frontal setae. Among the re-
maining 13 d paratypes, nine (19%) had
three and four frontal setae on different
sides of their heads; two d¢ (4%) each had
two and three frontal setae, and one d each
had three and five or four and five frontal
setae located asymmetrically.
Regarding the wings, the relationship of
the subapical and apical bands showed vari-
ation. The holotype and 48 @ paratypes
(69%) of a total of 71 @ types with intact
wings had the subapical and apical bands
fully separated (Fig. 2A); 13 2 paratypes
(18%) had the subapical and apical bands
Us:
narrowly separated; and in the remaining 9
2 (13%), these bands were narrowly or
widely joined in cell r, in eight 2 and one
2, respectively. The allotype and 21 ¢d
paratypes (45%) of a total of 49 d types
with intact, fully formed wings had the sub-
apical and apical bands fully separated (Fig.
2B); 19 ¢ paratypes (39%) had these bands
narrowly separated in at least one wing (in
the other wing in four d, these bands were
narrowly joined in cell r, or r,,,); and in the
remaining eight d paratypes (16%), these
bands were broadly joined in four 6 or nar-
rowly joined in four ¢ in cell r, or r5,3.
In the holotype and 35 @& paratypes
(51%) of the same 71 @ types with fully
developed wings, the discal band extended
across vein A, + CuA, into the anal lobe
in both wings (Fig. 2A); in 27 2 paratypes
(37%), this band ended at this vein in both
wings; in two 2 paratypes (3%) each wing
showed a different one of these two char-
acters. In only three (4%) of the remaining
? paratypes, the discal band ended before
and did not reach vein A, + CuA,. In the
allotype and 39 ¢d paratypes (80%) of the
same 49 ¢ types with intact, fully formed
wings, the discal band ended before and did
not touch vem Ay -- CuwA> (Fics 2B): or
reached this vein in nine (18%) other 3
paratypes, but crossed this vein into the
anal lobe in only one d paratype. These
data, therefore, provide the first document-
ed incidence of sexual dimorphism in the
wing pattern of a species of Procecido-
chares (Foote et al. 1993); although, this
dimorphism pales in comparison with wing
pattern sexual dimorphism recently report-
ed by us for certain species of Aciurina
(Goeden and Teerink 1996a, b, c), or as
long known for certain Trupanea spp.
(Foote, ctjala 1993):
Diagnosis.—The incomplete discal band
distinguishes P. lisae from all previously
described species of Procecidochares. Two
other characters distinguishing the adults of
P. kristineae and P. lisae from other Pro-
cecidochares spp. are the pterostigma being
lighter brown in the basal fifth to half than
74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
in the apical part, in combination with one
pair of orbital setae. These species can then
be readily separated on the basis of their
wing patterns and the color of the genal
spot as described in the partial key provided
above. The pterostigmal character is shared
only by P. flavipes, P. kristineae, and P.
lisae, with P. flavipes separated by its pos-
session of two pairs of orbital setae. The
basal and discal bands are joined only in
the wings of P. kristineae, P. lisae, and P.
stonei, with P. lisae readily distinguished
by the broad anterior juncture of these
bands and by the shortened discal bands.
Types.—Holotype, 2; Mountain Pass,
1430-m elevation, NE San Bernardino Co.,
California; 30.iv.1981; RDG, coll.; reared
from axillary bud gall on A. eriocentra
(USNM); allotype, 6, same data as holo-
type (USNM). Paratypes: CALIFORNIA:
17 3 and 17 92; same data as holotype (5
6 and5 ¢ to USNM, 6 6 and 6 2 to CAS).
5 6 and 1 2,2 6 and 2 2; same location
data as holotype; 19.iv.1971, 26.vi.1971, re-
spectively, RDG & DWR, coll. 5 3 and 3
2,2 2,106 and 11 2; same location data
as ‘holotype; 9:-v- 1991. 9:v.1992. 9.v. 1993;
respectively; RDG & JAT, coll. 8 ¢ and 15
?; Halloran Springs, NE San Bernardino
Go 191iv.1971; RDG PWR colly 372:
Halloran Springs, NE San Bernardino Co.;
291v. 1981) RDGcoll” 12'S and ie es
Hackberry Mountain, NE San Bernardino
Co-; 220% 1970.4 111-197 1 respectively:
RDG & DWR, coll. 3 3d and 4 @; Granite
Cove, NW San Bernardino Co., 25.v.1971;
RDG & DWR, coll. (Remaining 36 d and
50 @ paratypes along with numerous swept
and/or damage-reared, voucher and non-
voucher specimens examined and identified
are held in research collection of RDG).
Etymology.—Procecidochares lisae is
named for my older daughter, Lisa Marie
Goeden, a unique woman of many accom-
plishments.
IMMATURE STAGES
Procecidochares kristineae.—Egg: Eggs
of P. kristineae (Fig. 3A, B), were de-
Fig. 3.
itus; (B) anterior end, aeropyles.
Egg of Procecidochares kristineae (A) hab-
scribed by Silverman and Goeden (1980).
Chorion smooth (Fig. 3A); pedicel with few
aeropyles (Fig. 3B).
Third instar: Superficially smooth, elon-
gate cylindrical, tapering anteriorly, lacking
minute acanthae (Fig. 4A); gnathocephalon
conical, broad dorsally, smooth with few
rugose pads (Fig. 4B); paired dorsal sen-
sory organs dorsomediad of anterior sen-
sory lobes, consisting of dome-shaped pa-
pilla (Fig. 4B-1, 4C-1); anterior sensory
lobes bear terminal sensory organ (Fig.
4C-2), pit sensory organ (Fig. 4C-3), lateral
sensory organ (Fig. 4C-4), and supralateral
sensory organ (Fig. 4C-5); stomal sense or-
gans ventrad of anterior sensory lobe (Fig.
4B-2); lateral sensillum ventrolaterad of
stomal sense organs (Fig. 4B-3); mouth
hooks tridentate, teeth conical, stout (Fig.
4B-4); median oral lobe fleshy, tapering an-
teriorly, attached to labial lobe (Fig. 4B-5);
labial lobe with two pore sensilla; ventro-
VOLUME 99, NUMBER 1
wo
the
; J
{sku xie0@ oo1e “f8.00
Fig. 4.
15KUY K2666 6616.
1a. au
ISKU 41166 8626
Third instar of Procecidochares kristineae: (A) habitus, anterior to left; (B) gnathocephalon, anterior
view, 1—dorsal sensory organ, 2—stomal sense organ, 3—lateral sensillum, 4—mouth hooks, 5—median oral
lobe, 6—ventrolateral sensillum; (C) anterior sensory lobe, 1—dorsal sensory organ, 2—terminal sensory organ,
3—pit sensory organ, 4—lateral sensory organ, 5—supralateral sensory organ; (D) anterior thoracic spiracles;
(E) second abdominal lateral spiracular complex, 1—spiracle, 2—verruciform sensillum; (F) posterior spiracular
plate, 1—rima, 2—interspiracular process.
lateral sensillum ventrolaterad of mouth lu-
men (Fig. 4B-6); prothorax smooth,
verruciform sensilla circumscribe dorsal
half of anterior margin, anterior thoracic
spiracles on posterior margin consist of
three ovoid papillae (Fig. 4D); meso- and
metathoracic lateral spiracular complexes
consist of an open spiracle; abdominal lat-
eral spiracular complex consist of an open
spiracle (Fig. 4E-1) and a single verruci-
form sensillum (Fig. 4E-2); caudal segment
bears posterior spiracular plates (Fig. 4F);
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
BSKY K2680
15KU x2688
@41i3
Fig. 5.
15KU ¥3660 6815
Second instar of Procecidochares kristineae: (A) gnathocephalon, anterior view, 1—dorsal sensory
organ, 2—stomal sense organ, 3—mouth hooks, 4—ventrolateral sensillum; (B) anterior sensory lobe, 1—dorsal
sensory organ, 2—terminal sensory organ, 3—lateral sensory organ, 4—supralateral sensory organ; (C) gnath-
ocephalon, anteriolateral view, 1—mouth hooks, 2—median oral lobe; (D) anterior thoracic spiracles; (E) pos-
terior spiracular plates, 1—rima, 2—interspiracular process; (F) compound sensillum, 1—verruciform sensillum,
2—-stelex sensillum.
posterior spiracular plates with three ovoid
rimae, ca. 0.024 mm in length (Fig. 4F-1),
and four spiniform interspiracular process-
es, longest measuring 0.01 mm in length
(Fig. 4F-2); compound sensilla ventrad of
posterior spiracular plates consist of a ver-
ruciform sensillum and a stelex sensillum.
Second instar: Superficially smooth, bar-
rel-shaped, rounded anteriorly and posteri-
orly; gnathocephalon conical, smooth with
few rugose pads (Fig 5A); paired dorsal
sensory organs consist of a dome-shaped
papilla (Fig. 5A-1, B-1); anterior sensory
lobe bears terminal sensory organ (Fig.
VOLUME 99, NUMBER |
15KY x8 6@ae
LT
Fig. 6. First instar of Procecidochares kristineae: (A) gnathocephalon, anteriolateral view, 1—dorsal sensory
organ, 2—mouth hooks; (B) anterior sensory lobe, 1—dorsal sensory organ, 2—terminal sensory organ, 3—pit
sensory organ, 4—lateral sensory organ, 5—supralateral sensory organ; (C) gnathocephalon, anteriolateral view,
1—mouth hooks, 2—median oral lobe; (D) caudal segment, posterior spiracular plates, 1—rima, 2—interspi-
racular process, 3—stelex sensillum.
5B-2), pit sensory organ, lateral sensory or-
gan (Fig. 5B-3), and supralateral sensory
organ (Fig. 5B-4); stomal sense organ ven-
trad of anterior sensory lobe (Fig. 5A-2);
mouth hooks tridentate, teeth conical, apical
tooth longest (Fig. SA-3, C-1); median oral
lobe laterally flattened, rounded apically
(Fig. SC-2); ventrolateral sensillum ventro-
laterad of mouth lumen (Fig. 5A-4); ante-
rior thoracic spiracles on posterior margin
of prothorax, consist of three ovoid papillae
(Fig. 5D); lateral spiracular complex not
observed; caudal segment bears posterior
spiracular plates (Fig. 5E); posterior spirac-
ular plates with three ovoid rimae, ca. 0.009
mm in length (Fig. SE-1) and four spini-
form interspiracular processes, longest mea-
suring 0.005 mm (Fig. 5E-2); compound
sensilla ventrad of posterior spiracular
plates, consist of a verruciform sensillum
(Fig. 5F-1), and a stelex sensillum (Fig.
5F-2).
First instar: Superficially smooth, barrel-
shaped, rounded anteriorly and posteriorly,
minute acanthae circumscribe segmental
lines; gnathocephalon conical, smooth,
lacking rugose pads (Fig. 6A); dorsal sen-
sory organ consists of a dome-shaped pa-
pilla (Fig. 6A-1, B-1); anterior sensory
lobes bear terminal sensory organ (Fig.
6B-2), pit sensory organ (Fig. 6B-3), lateral
sensory organ (Fig. 6B-4), and supralateral
sensory organ (Fig. 6B-5); stomal sense or-
gans indistinct; mouth hooks bidentate, api-
cal tooth with v-shaped frontal groove
along entire length (Fig. 6A-2, 6C-1); me-
dian oral lobe rounded apically (Fig. 6C-2);
anterior thoracic spiracles absent; lateral
78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
spiracular complexes not observed; caudal
segment bears posterior spiracular plates
(Fig. 6D); posterior spiracular plates with
two ovoid rimae, ca. 0.002 mm (Fig. 6D-1),
and four ovoid interspiracular processes,
longest measuring 0.001 mm (Fig. 6D-2);
stelex sensillum ventrad of posterior spirac-
ular plates (Fig. 6D-3).
Puparium: Superficially smooth, elon-
gate-ellipsoidal (Fig. 7A); anterior end
bears invagination scar (Fig. 7B-1), and an-
terior thoracic spiracles (Fig. 7B-2); caudal
segment bears posterior spiracular plates
(Fig. 7C); posterior spiracular plates with
three ovoid rimae, ca. 0.03 mm in length
(Fig. 7C-1), and four spiniform interspirac-
ular processes, longest measuring 0.01 mm
(Fig. 7C-2). Measurements of P. kristineae
puparia were provided by Silverman and
Goeden (1980).
Procecidochares lisae.—Egg: Fifty ova
of P. lisae were white, opaque, smooth;
with an elongate-ellipsoidal body, 0.57 +
0.005 (range, 0.52—0.64) mm long, 0.17 +
0.002 (range, 0.16—0.20) mm wide, smooth-
ly rounded at tapered posterior end; and
with a short, peg-like anterior pedicel, 0.02
mm long by 0.03 wide.
Third instar: Superficially smooth, elon-
gate cylindrical, tapering anteriorly, lacking
minute acanthae (Fig. 8A); gnathocephalon
conical, broad dorsally, smooth with few
rugose pads (Fig. 8B); paired dorsal sen-
sory organs dorsomediad of anterior sen-
sory lobes, consisting of dome-shaped pa-
pilla (Fig. 8B-1); anterior sensory lobes
bear terminal sensory organ (Fig. 8B-2), pit
sensory organ (Fig. 8B-3), lateral sensory
organ (Fig. 8B-4), and supralateral sensory
organ (Fig. 8B-5); stomal sense organs ven-
trad of anterior sensory lobe; lateral sensil-
lum ventrolaterad of stomal sense organs
(Fig. 8B-6); mouth hooks tridentate, apical
tooth longest (Fig. 8C-1); median oral lobe
fleshy, laterally flattened, tapering anterior-
ly, (Fig. 8C-2); ventrolateral sensillum ven-
trolaterad of mouth lumen (Fig. 8C-3); pro-
thorax smooth, verruciform sensilla circum-
scribe dorsal half of anterior margin; ante-
; a0. BB
; ~ ype A
9024 160. 6U Cc
“TSKY ¥320
Fig. 7. Puparium of Procecidochares kristineae:
(A) habitus, anterior to left; (B) anterior end, 1—in-
vagination scar, 2—anterior thoracic spiracles; (C) cau-
dal segment, posterior spiracular plates, |—rima, 2—
interspiracular process.
rior thoracic spiracles consist of three ovoid
papillae (Fig. 8D); meso- and metathoracic
lateral spiracular complexes consist of an
open spiracle; abdominal lateral spiracular
complex consist of an open spiracle (Fig.
8E-1) and a single verruciform sensillum
(Fig. 8E-2); caudal segment bears posterior
VOLUME 99, NUMBER |
15K A,
\
b aid
15KU.x1600 ° ‘9019
Fig. 8.
i9. au E
Third instar of Procecidochares lisae: (A) habitus, anterior to left; (B) gnathocephalon, anteriolateral
79
@aa4 \ 16. aU
SKY K1146
view, 1—dorsal sensory organ, 2—terminal sensory organ, 3—pit sensory organ, 4—lateral sensory organ, 5—
supralateral sensory organ, 6—lateral sensillum; (C) gnathocephalon, anterior view, |—mouth hooks, 2—median
oral lobe, 3—ventrolateral sensillum; (D) anterior thoracic spiracles; (E) first abdominal lateral spiracular com-
plex, 1—spiracle, 2—verruciform sensillum; (F) posterior spiracular plate, 1—rima, 2—interspiracular process.
spiracular plates; posterior spiracular plates
with three ovoid rimae, ca. 0.025 mm in
length (Fig. 8F-1), and four spiniform in-
terspiracular processes, longest measuring
0.01 mm in length (Fig. 8F-2); compound
sensilla ventrad of posterior spiracular
plates consist of two verruciform sensilla.
Second instar: Superficially smooth, cy-
lindrical, rounded anteriorly and posterior-
ly; gnathocephalon conical, smooth with
few rugose pads; paired dorsal sensory or-
gans consist of a dome-shaped papilla (Fig.
9A-1); anterior sensory lobe bear terminal
sensory organ (Fig. 9A-2), pit sensory or-
80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
aga2°7 18.8
\
15SkKU X240o
Fig. 9.
Second instar of Procecidochares lisae: (A) gnathocephalon, anterior view, 1—dorsal sensory organ,
2—terminal sensory organ, 3—pit sensory organ, 4—lateral sensory organ, 5—supralateral sensory organ; (B)
gnathocephalon, anteriolateral view, 1—stomal sense organ, 2—median oral lobe, 3—mouth hooks; (C) anterior
thoracic spiracles; (D) caudal segment, posterior spiracular plates, 1—rima, 2—interspiracular process.
gan (Fig. 9A-3), lateral sensory organ (Fig.
9A-4), and supralateral sensory organ (Fig.
9A-5); stomal sense organ ventrad of an-
terior sensory lobe (Fig. 9B-1); mouth
hooks tridentate, teeth conical (Fig. 9B-3);
median oral lobe fleshy, laterally flattened
(Fig. 9B-2); anterior thoracic spiracles on
posterior margin of prothorax, consist of
two-three ovoid papillae (Fig. 9C); lateral
spiracular complex not observed; caudal
segment bears posterior spiracular plates;
posterior spiracular plates with three ovoid
rimae, ca. 0.009 mm in length (Fig. 9D-1)
and four spiniform interspiracular process-
es, longest measuring 0.004 mm (Fig.
9D-2); compound sensilla ventrad of pos-
terior spiracular plates were obscured in
prepared specimens.
First instar: Superficially smooth, cylin-
drical, tapering anteriorly, rounded posteri-
orly, minute acanthae circumscribe segmen-
tal lines (Fig. 10A); gnathocephalon coni-
cal, smooth, lacking rugose pads (Fig.
10B); dorsal sensory organs large,
dome-shaped (Fig. 1OB-1, C-1); anterior
sensory lobes bear terminal sensory organ
(Fig. 10C-2), pit sensory organ (Fig. 10C-
3), lateral sensory organ (Fig. 10C-4), and
supralateral sensory organ (Fig. 10C-5);
stomal sense organs indistinct; mouth hooks
bidentate, apical tooth with v-shaped frontal
groove along midline (Fig. 1OB-2, D-1);
ventrolateral sensillum ventrolaterad of
mouth lumen (Fig. 1OB-3, D-2); anterior
thoracic spiracles absent; lateral spiracular
complexes not observed; caudal segment
bears posterior spiracular plates (Fig. 10E-
1); posterior spiracular plates with two
ovoid rimae, ca. 0.002 mm in length (Fig.
10F-1), and four, very reduced spiniform
VOLUME 99, NUMBER |
15KY
$1
Fig. 10. First instar of Procecidochares lisae: (A) habitus, anterior to left; (B) gnathocephalon, lateral view,
1—dorsal sensory organ, 2—mouth hooks, 3—ventrolateral sensillum; (C) gnathocephalon, 1—dorsal sensory
organ, 2—terminal sensory organ, 3—pit sensory organ, 4
lateral sensory organ, 5—supralateral sensory organ;
(D) gnathocephalon, ventral view, 1—mouth hooks, 2—ventrolateral sensillum; (E) caudal segment, 1—posterior
spiracular plates, 2—verruciform sensillum, 3—compound sensillum, verruciform sensillum; (F) posterior spi-
racular plates, 1—rima, 2—interspiracular process.
interspiracular processes, longest measuring
0.001 mm (Fig. 1OF-2); two verruciform
sensilla dorsolaterad of posterior spiracular
plates (Fig. 1|OE-2); compound sensilla ven-
trad of posterior spiracular plates consist of
two verruciform sensilla (Fig. 1OE-3).
Puparium: Superficially smooth, elon-
gate-ellipsoidal (Fig. I1A); anterior end
bears invagination scar (Fig. 11B-1), and
anterior thoracic spiracles (Fig. 11B-2);
caudal segment bears posterior spiracular
plates, each with three ovoid rimae, ca. 0.04
82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
15KV
Ace
9
6665 166. 68U | &
ISKY K266
Fig. 11. Puparium of Procecidochares lisae: (A)
habitus, anterior to left; (B) anterior end, 1—invagi-
nation scar, 2—anterior thoracic spiracle; (C) caudal
segment, 1—rima, 2—interspiracular process.
mm in length (Fig. 11C-1), and four thorn-
like interspiracular processes, longest mea-
suring 0.007 mm (Fig. 11C-2). Fifty-six pu-
paria averaged 4.01 + 0.05 (range, 3.24-
4.60) mm in length; 1.83 + 0.03 (range,
1.40-2.35) mm in width.
Procecidochares kristineae and P. lisae
larvae are morphologically very similar.
The only difference noted was the com-
pound sensilla ventrad of the posterior spi-
racular plates; in P. kristineae there was a
verruciform sensillum and a stelex sensil-
lum, but both sensilla in P. lisae were ver-
ruciform. Thus, morphological differentia-
tion between these two, possibly sibling
species on separate, closely related hosts is
minimal between their larvae, but distinc-
tive between their adults. Similar morpho-
logical differences between larvae and
adults of two sympatric, nongallicolous,
also probably sibling species, Trupanea ni-
gricornis and T. bisetosa recently was re-
ported (Knio et. al. 1996).
In turn, Procecidochares kristineae and
P. lisae are very similar to P. stonei (Green
et al. 1993). The noticeable difference be-
tween P. stonei and these two species in the
third instar larva is that in the lateral spi-
racular complex, the verruciform sensillum
is anteriorad of the spiracle in P. stonei not
posteriorad. In the first instar of P. stonei,
the mouth hooks are not grooved dorsally
and the interspiracular processes are multi
branched and blade-like (Green et al. 1993),
not ovoid or spiniform and unbranched as
in P. kristineae and P. lisae, respectively.
Procecidochares kristineae and P. lisae are
the first species of Tephritidae described
with a v-shaped frontal groove in the apical
tooth (Goeden and Headrick 1992; Goeden
and Teerink 1996a, b, c; Goeden et al.
1994b; Green et al. 1993; Headrick and
Goeden 1990; Headrick et al. 1995; Knio
et al. 1996). The nongallicolous, flower
head infesting species P. flavipes differs
from all three of the above gall-forming
species in having thoracic segments cov-
ered with small, polygonal rugose pads and
the ventral intersegmental areas and dorsum
of the caudal segment bearing minute acan-
thae (Goeden et al. 1994).
DISTRIBUTION AND Hosts
Both Procecidochares kristineae and P.
lisae are true monophages, each forming
galls only on its sole host-plant species,
Ambrosia dumosa and A. eriocentra, re-
spectively. Procecidochares kristineae is
VOLUME 99, NUMBER 1
widely associated with A. dumosa in the
Colorado and Mojave deserts of southern
California (Goeden and Ricker 1976a, Sil-
verman and Goeden 1980). Accordingly,
the geographic distribution of P. kristineae
may coincide wholly or partly with that of
A. dumosa in southern Nevada, southwest-
ern Utah, and western and southwestern Ar-
izona (Benson and Darrow 1981), and
southward into the Sonoran Desert of Baja
California and Sonora, Mexico (Shreve and
Wiggens 1964). The distributions of P. li-
sae and A. eriocentra in southern California
are restricted to a small part of the north-
eastern Mojave Desert (Goeden and Ricker
1976b), but probably coincides wholly or
partly with the wider distribution of this na-
tive ragweed in southern Nevada, south-
western Utah, and northwestern and central
Arizona (Benson and Darrow 1981).
Estimating tephritid distribution from
host-plant distribution is problematic be-
cause some gall-forming tephritid flies are
known to be less widely distributed than
their hosts, e.g., A. bigeloviae (Cockerell)
and A. trixa Curran are widely allopatric,
sympatric, or absent on Chrysothamnus
nauseosus (Pallas) Britton in different parts
of Utah, Arizona, New Mexico, and Colo-
rado (Dodson and George 1986). Similarly,
A. trixa, but apparently not A. bigeloviae,
occurs in California on C. nauseosus which
is a widespread plant species in southern
California (D. H. Headrick, RDG, and JAT,
unpublished data).
BIOLOGY
The biology of P. kristineae was de-
scribed in considerable detail by Silverman
and Goeden (1980). The following account
largely concerns P. lisae, which then serves
as a basis for comparison with P. kristineae
and two other taxonomically-close Proce-
cidochares spp. studied previously in south-
ern California, P. stonei (Green et al. 1993)
and P. flavipes (Goeden et al. 1994a).
Egg.—Most eggs are inserted singly and
basally in axillary buds of the current sea-
son’s branch growth. Some buds held two
83
or rarely three eggs, presumably oviposited
by different females, judging by the slightly
different degrees of development of two or
three first instars found associated with sin-
gle buds in field-collected branch samples
(see below). Two eggs of P. kristineae per
axillary bud also were reported as rare by
Silverman and Goeden (1980). Adjacent
axillary buds of infested branches bore eggs
probably deposited in sequence by individ-
ual females as they moved in sequence to
the next more-proximal axils, as described
for P. kristineae (Silverman and Goeden
1980).
Larva.—Eggs hatch about a week after
their deposition (Goeden and Silverman
1980, Green et al. 1993), but then the first
instar larvae persist as the longest-lived
stage in the life cycle of P. lisae within tiny
open cavities of incipient galls (Fig. 12A).
Single, aestivating first instars were found
Overwintering in ovoidal cavities 0.74 +
0.03 (range, 0.54—0.93) mm long by 0.54
+ 0.02 (range 0.37—0.74) mm wide formed
within, beneath, or immediately laterad of
26 axillary buds. Four pairs each of these
first instars were associated with single ax-
illary buds, and three first instars with an-
other bud, documenting again multiple ovi-
positions in single buds. Each of these lar-
vae continued to develop separately, as all
galls examined of P. lisae, like P. kristineae
(Silverman and Goeden 1980), contained
only one larva in each. Thus, sometimes
two or rarely three unilocular galls of P.
lisae arose at the same axil; although, only
one gall per axil was usual for both P. kris-
tineae and P. lisae (Silverman and Goeden
1980). However, larvae of P. stonei develop
gregariously in unilocular galls (Green et al.
1995);
The aforementioned 26 galls were sam-
pled on 7.1i1.1995, so that not only were
these aestivating first instars the overwin-
tering stage of P. lisae, but also the stage
within which this species had persisted for
6 or 7 months since the previous April or
May, when their parent adults emerged,
mated, and oviposited. Both P. kristineae
84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
i” ec
Fig. 12. Life stages of Procecidochares lisae on Ambrosia eriocentra: (A) axillary bud gall containing
aestivating first instar, (B) sagittal section of preceding axillary bud gall, (C) axillary bud gall containing second
instar, (D) sagittal section of gall containing second instar, (E) full-size gall, (F) sagittal section through full-
size gall containing a puparium, and showing exit channel excavated by late third instar, (G) lateral view of
male, (H) lateral view of gravid female, (I) mating pair, dorsal view. Lines = 1 mm.
VOLUME 99, NUMBER 1
and P. stonei also overwinter as first instars
in incipient galls (Silverman and Goeden
1980, Green et al. 1993). The tiny, over-
wintering galls of P. lisae were undetecta-
ble in the field (Fig. 12A, B); although
those of P. kristineae were readily dis-
cerned (Silverman and Goeden 1980). The
former galls caused only a slight axillary
swelling along the leafless branches of dor-
mant A. eriocentra, unlike overwintering
galls of P. kristineae on A. dumosa which
incorporated juvenile leaves (Silverman and
Goeden 1980). The axils on branches where
incipient galls of P. lisae were borne aver-
aged 3.04 + 0.05 (range, 2.5—3.6) mm in
diameter (n = 21) and were located an av-
erage of 15.8 + 1.1 (range, 6—24.5) cm
from branch apex.
Following substantial late-winter/early-
spring rainfall, host plants, galls, and larvae
of P. lisae concurrently resume growth as
also reported for P. kristineae (Silverman
and Goeden 1980) and P. stonei (Green et
al. 1993). Twenty-five galls of P. lisae con-
taining second instars (Fig. 12C, D) aver-
aged 5.72 + 0.35 (range, 2.41—11.40) mm
in length; 5.44 + 0.34 (range, 2.28—11.40)
mm in width. These spheroidal galls (Fig.
12C) contained single, ovoidal to spheroi-
dal cavities (Fig. 12D) that averaged 2.53
* 0.21 (range, 1.14—-5.70) mm in length
and 1.97 + 0.16 (range, 0.57—4.56) mm in
width. The gall walls averaged 1.12 + 0.10
(range, 0.42—2.85) mm in thickness.
The larvae of P. lisae quickly grew and
molted into the third and last larval instar
within mostly spheroidal galls (Fig. 12E,
F), 21 of which averaged 8.0 + 0.4 (range,
4.3—11) mm in length and 7.1 + 0.3 (range,
4.2-10) mm in width. The central ovoidal
or spheroidal cavities were enlarged by lar-
val feeding to an average of 4.1 + 0.2
(range, 1.42—5.13) mm in length and 3.17
+ 0.18 (range, 1.42—5.13) mm in width.
The lateral wall thickness averaged 1.18 +
0.04 (range, 0.57—-2.15) mm. Prior to pu-
pariation, the third instar chews an exit tun-
nel through the gall wall (Fig. 12F) ending
in a thin, circular to oval, epidermal win-
85
dow 1.4 + 0.04 (range, 1.2—1.7) mm (n =
13) in diameter.
Pupa.—Fifty-six galls containing puparia
(Fig. 12F), and thus of maximum size and
final subspheroidal shape, averaged 10.2 +
0.4 (range, 3.9—16) mm in length; 8.9 + 0.3
(range, 3.5—16) mm in width. The ovoidal
to spheroidal cavities of these mature galls
averaged 4.8 + 0.2 (range, 3.1—3.4) mm in
length; 4.0 + 0.1 (range, 2.6—6.2) mm in
width. The lateral walls of these galls av-
eraged 1.58 + 0.06 (range, 0.51—2.58) mm
in thickness.
Adult.—The adults (Fig. 12G, H) of P.
lisae that emerged from excised, field-col-
lected galls were sexually mature, or nearly
so, each female containing many full-size
ova. These proovigenic females mated in
laboratory cagings from | to 7 days-old (n
= 59). (ries 21). Jen aales, lived, Jed) =
0.8 (range, 7-16) days; 11 nonovipositing
females, 8.7 + 0.7 (range, 5-12) days in
insectary cagings. These mean longevities
fell between the 1l-week averages for P.
kristineae (Silverman and Goeden 1980)
and the 3-week averages for P. stonei
(Green et al. 1993) under similar insectary
conditions. This compares with longevities
that averaged 42 days for males and 39 and
102 days for two series of synovigenic fe-
males (n = 11 and 20, respectively) of P.
flavipes (Goeden et al. 1994a), which has
an aggregative life cycle different from the
circumnatal life cycles of P. kristineae, P.
lisae, and P. stonei (Headrick and Goeden
1994).
The reproductive behavior of P. lisae re-
sembled that of P. kristineae described by
Silverman and Goeden (1980) and by D. H.
Headrick from field observations (see be-
low), and that of P. stonei described in
greater detail by Green et al. (1993). Fur-
thermore, a lack of courtship behavior and
exclusive exhibition of enantion type of
wing movements by both sexes reported by
Headrick and Goeden (1994) as character-
istic of circumnatal species of Procecido-
chares also was confirmed for P. lisae.
Wing enantion as defined and described by
86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Green et al. (1993) and Headrick and Goe-
den (1994) was observed for P. lisae as
males tracked females prior to copulation,
during copulatory induction behavior, and
following postcopulatory separation. Cop-
ulation by only one pair of P. lisae was
observed from start to finish for 62 min, but
nine more partially observed matings all
were of shorter duration. Disengagement by
three different pairs of adults was observed,
whereby the males turned and walked off
and away from the females as they concur-
rently pulled their genitalia from the fe-
males’ cloacas in one continuous motion in
15, 18, and 30 sec, respectively. The wings
of both sexes of P. lisae usually were parted
at 45° and centered over their midlines dur-
ing copulation; whereas, Silverman and
Goeden (1980) reported that males of P.
kristineae usually were only slightly parted
during copulation, a position sometimes
also adopted by P. lisae males.
Field observations.—Field observations
of P. kristineae behaviors on Ambrosia du-
mosa were made by D. H. Headrick at
Snow Creek, Riverside Co., 2-9 April
1992. Most observations were made for 1—
3 h daily on three large shrubs all bearing
newly formed, immature inflorescences.
The oviposition behavior of P. kristineae
females was highly stereotypical. Each fe-
male began by walking from the base of a
current season’s branch to the base of the
raceme, where she turned head-downward,
recurved her abdomen and oviscape, and
probed with the apex of the aculeus into a
leaf axil. After ovipositing or probing, the
female moved basally to the next leaf axil
and probed or oviposited in the same man-
ner. When the female reached the base of
the new growth, she moved to another
branch and repeated the above behaviors.
Females probed many leaf axils, but ovi-
posited into only 1 to 4 axils per branch.
Oviposition times averaged 2 min 9 sec (n
= 4; range, 2 min—2 min 20 sec). Females
frequently stopped to rest and groom be-
tween Ovipositions.
Mating behavior of P. kristineae was
similar to that reported by Silverman and
Goeden (1980). Males encountered females
either while perched on top of branches or
while moving through the crown. Males
tracked females slowly from behind. Males
sat still in a passive position if the female
turned to face him, or paused nearby. Males
mounted females by jumping onto their
dorsa, either head first or from behind.
Once mounted, males clasped the costal
margins of the female’s wings near the base
with the foretarsal claws; the middle legs
were wrapped around the middle of the ab-
domen of the female and the hind legs were
bent and crossed underneath the abdomen
of the female. Once mounted, males im-
mediately began rubbing their hind legs
asynchronously anterior to posterior under-
neath the abdomen of the female. Simulta-
neously, the abdomen of the female was
raised and the apex of the ovipositor was
placed into the epandrium; exsertion of the
aculeus followed (n = 5). Copulation in the
field lasted 30 min (1 = 2).
Seasonal history.—Procecidochares li-
sae usually is univoltine, but occasionally,
following substantial late-summer rainfall
stimulating renewed host-plant growth, pro-
duces a facultative second generation. Pro-
cecidochares kristineae is normally bivol-
tine throughout its much wider range in
southern California, but as reported by Sil-
verman and Goeden (1980) can _ locally
show univoltine, biennial, and even trivol-
tine reproduction (unpublished data), de-
pending on whether rainfall locally triggers
resumption of gall and larval development
concurrent with host-plant regrowth. Both
species, along with P. stonei, exhibit the
circumnatal life cycle defined and described
by Headrick and Goeden (1994).
Natural enemies.—The following chal-
cidoid Hymenoptera were identified as par-
asitoids of P. lisae: Eurytoma sp. and E.
veronia Bugbee (Eurytomidae), as primary,
solitary parasitoids reared from mature
galls; Halticoptera sp. (Pteromalidae), as a
primary, internal, larval-pupal parasitoid;
Tetrastichus sp. (Eulophidae), as a gregar-
VOLUME 99, NUMBER 1
ious, puparial endoparasitoid; Torymus sp.
(Bedeguaris group), Torymus sp. (Tubicola
group), 7. capillaceus (Hiiber) (Torymi-
dae), probably as primary, solitary, larval
ectoparasitoids as reared from immature
galls.
Silverman and Goeden (1980) reported
10 species of parasitic Hymenoptera reared
from fully formed galls of P. kristineae, an
unidentified vertebrate predator of the lar-
vae and identified spider predator of the
adults, and two species of inquiline weevils.
Green et al. (1993) subsequently reported
four species of primary chalcidoid parasit-
oids individually reared from puparia of P.
stonei as well as a hyperparasitized larval
parasitoid. The genera of parasitoids com-
mon to all three Procecidochares hosts
were Eurytoma, Halticoptera, and Tetras-
tichus.
ACKNOWLEDGMENTS
We sincerely thank A. C. Sanders, Cu-
rator of the Herbarium, Department of Bot-
any and Plant Sciences, University of Cal-
ifornia, Riverside, for identification of
plants. The parasitoids variously were iden-
tified by H. E. Andersen, Huntington
Beach, CA; G. Gordh, now with the De-
partment of Entomology, University of
Queensland, Australia; or J. Luhman, now
with the Minnesota Department of Agricul-
ture, St. Paul, MN. We also are grateful to
FL. Blanc, D. H. Headrick, A. L. Norrbom,
J. D. Pinto, and G. J. Steck for their helpful
comments on earlier drafts of this paper.
LITERATURE CITED
Benson, L. and R. A. Darrow. 1981. Trees and Shrubs
of the Southwestern Deserts. University of Ari-
zona Press, Tucson.
Blanc, FE L. and R. H. Foote. 1961. A new genus and
five new species of California Tephritidae. Pan-
Pacific Entomologist 37: 73-83.
Dodson, G. and S. B. George. 1986. Examination of
two morphs of gall-forming Aciurina (Diptera: Te-
phritidae). Ecological and genetic evidence for
species. Biological Journal of the Linnean Society
29: 63-79.
Foote, R. H., EF L. Blanc, and A. L. Norrbom. 1993.
Handbook of the Fruit Flies (Diptera: Tephritidae)
87
of America North of Mexico. Cornell University
Press, Ithaca, New York.
Goeden, R. D. and D. H. Headrick. 1992. Life history
and descriptions of immature stages of Neaspilota
viridescens Quisenberry (Diptera: Tephritidae) on
native Asteraceae in southern California. Proceed-
ings of the Entomological Society of Washington
94: 59-77.
Goeden, R. D., D. H. Headrick, and J. A. Teerink.
1994a. Life History and descriptions of immature
stages of Procecidochares flavipes Aldrich (Dip-
tera: Tephritidae) on Brickellia spp. in southern
California. Proceedings of the Entomological So-
ciety of Washington 96: 288—300.
1994b. Life history and description of im-
mature stages of Paroxyna genalis (Thomson)
(Diptera: Tephritidae) on native Asteraceae in
southern California. Proceedings of the Entomo-
logical Society of Washington 96: 612-629.
1995a. Life history and descriptions of im-
mature stages of Valentibulla californica (Coquil-
lett) (Diptera: Tephritidae) on Chrysothamnus
nauseosus (Pallas) Britton in southern California.
Proceedings of the Entomological Society of
Washington 97: 548-560.
1995b. Life history and description of im-
mature stages of Urophora timberlakei Blanc and
Foote (Diptera: Tephritidae) on native Asteraceae
in southern California. Proceedings of the Ento-
mological Society of Washington 97: 780-791.
Goeden, R. D. and D. W. Ricker. 1976a. The phy-
tophagous insect fauna of the ragweed, Ambrosia
dumosa, in southern California. Environmental
Entomology 5: 45—SO0.
1976b. The phytophagous insect faunas of
the ragweeds, Ambrosia chenopodiifolia, A. eri-
ocentra, and A. ilicifolia. Environmental Ento-
mology 5: 923-930.
Goeden, R. D. and J. A. Teerink. 1996a. Life histories
and descriptions of adults and immature stages of
two cryptic species, Aciurina ferrugenia (Doane)
and A. michaeli new species (Diptera: Tephritidae)
on Chrysothamnus viscidiflorus (Hooker) Nuttall
in southern California. Proceedings of the Ento-
mological Society of Washington 98: 415—438.
. 1996b. Life history and descriptions of adults
and immature stages of Aciurina idahoensis
Steyskal (Diptera: Tephritidae) on Chrysothamnus
viscidiflorus (Hooker) Nuttall in southern Califor-
nia. Proceedings of the Entomological Society of
Washington 98: 681—694.
. 1996c. Life history and descriptions of adults
and immature stages of Aciurina semilucida
(Bates) (Diptera: Tephritidae) on Chrysothamnus
viscidiflorus (Hooker) Nuttall in southern Califor-
nia. Proceedings of the Entomological Society of
Washington 98: 752-766.
Green, J. EF, D. H. Headrick, and R. D. Goeden. 1993.
88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Life history and description of immature stages of
Procecidochares stonei Blanc & Foote on Vigui-
era spp. in southern California (Diptera: Tephrit-
idae). Pan-Pacific Entomologist 69: 18-32.
Headrick, D. H. and R. D. Goeden. 1990. Description
of the immature stages of Paracantha gentilis
(Diptera: Tephritidae). Annals of the Entomolog-
ical Society of America 83: 220-229.
1991. Life history of Trupanea californica
Malloch (Diptera: Tephritidae) on Gnaphalium
spp. in southern California. Proceedings of the En-
tomological Society of Washington 93: 559-570.
. 1993. Life history and description of imma-
ture stages of Aciurina thoracica (Diptera: Te-
phritidae) on Baccharis sarothroides in southern
California. Annals of the Entomological Society
of America 86: 68-79.
Headrick, D. H., R. D. Goeden, and J. A. Teerink.
1995. Life history and description of immature
stages of Euaresta stigmatica Coquillett (Diptera:
Tephritidae) on Ambrosia spp. (Asteraceae) in
southern California. Annals of the Entomological
Society of America 88: 58-71.
Knio, K. M., R. D. Goeden, and D. H. Headrick. 1996.
Descriptions of immature stages of Trupanea ni-
gricornis and T. bisetosa (Diptera: Tephritidae)
from southern California. Annals of the Entomo-
logical Society of America 89: 1-11.
Munz, P. A. 1974. A Flora of Southern California.
University of California Press, Berkeley.
Shreve, FE and I. L. Wiggens. 1964. Vegetation and
Flora of the Sonoran Desert, Vol. 2. Stanford Uni-
versity Press, Palo Alto, California.
Silverman, J. and R. D. Goeden. 1980. Life history
of a fruit fly, Procecidochares sp., on the ragweed
Ambrosia dumosa, (Gray) Payne, in southern Cal-
ifornia (Diptera: Tephritidae). Pan-Pacific Ento-
mologist 56: 283-288.
Wasbauer, M. W. 1972. An annotated host catalog of
the fruit flies of America north of Mexico (Dip-
tera: Tephritidae). California Department of Ag-
riculture, Bureau of Entomology, Occasional Pa-
pers 19: 1-72.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 89-100
LIFE HISTORY AND LABORATORY REARING OF OEDANCALA DORSALIS
(SAY) (HETEROPTERA: LYGAEIDAE), WITH DESCRIPTIONS OF
IMMATURE STAGES'!
LANE J. LOYA AND J. E. MCPHERSON
Department of Zoology, Southern Illinois University, Carbondale, IL 62901, U.S.A.
Abstract.—The life history of the seed bug Oedancala dorsalis (Say) was investigated
in southern [Illinois from June 1994—November 1995, and the immature stages were de-
scribed. The bug also was reared from egg to adult under controlled laboratory conditions.
This apparently bivoltine species fed and reproduced on Carex spp. Adults overwintered
and became active in April. Eggs were found from mid-May to early September in the
spikes of Carex crinita. Seasonal occurrences of adults and nymphs are discussed. Adults
were last observed in mid-October. This species was reared on C. crinita in the laboratory
at 26 + 0.5°C under a 16L:8D photoperiod. The incubation period averaged 13.32 days.
The five nymphal stadia averaged 6.34, 5.28, 4.52, 4.02, and 5.52 days, respectively.
Key Words:
Seed bug, Lygaeidae, southern Illinois, life history, laboratory rearing, im-
mature stages, descriptions, Carex
The family Lygaeidae contains approxi-
mately 4050 species worldwide (Slater and
O’Donnell 1995). Next to the family Miri-
dae, it is the second largest family of Het-
eroptera in America north of Mexico, where
it is represented by approximately 320 spe-
cies (Ashlock and Slater 1988). Lygaeids
are commonly called seed bugs, referring to
their habit of feeding on dry, mature seeds
(Sweet 1960).
In the United States and Canada, the ly-
gaeid subfamily Pachygronthinae contains
the three genera Oedancala Amyot and Ser-
ville, Pachygrontha Germar, and Phlegyas
Stal (Ashlock and Slater 1988). Only five
species of Oedancala occur within this geo-
graphic area (Ashlock and Slater 1988, Bar-
anowski and Slater 1989). Of these, O. dor-
salis Say is the most widespread, ranging
' Part of a thesis submitted to Southern Illinois Uni-
versity at Carbondale by the senior author in partial
fulfillment of the requirements for the M.S. degree in
zoology.
from Ontario, Quebec, and Maine south to
Florida, and west to South Dakota, Colo-
rado, and Texas, and is the only species of
the genus that has been reported from Illi-
nois (Ashlock and Slater 1988, Baranowski
and Slater 1989).
Oedancala dorsalis has been little stud-
ied. It has been swept from herbage in wet
meadows, pastures, and waste places
(Blatchley 1926), and reported to feed on
Carex, Cyperus (Van Duzee 1888, Slater
1951), Ceanothus, and marsh grasses (Slat-
er 1951). Van Duzee (1888) found adults
from May to September “‘about” Buffalo,
New York. Uhler (1887) stated it rarely was
found later than early summer in eastern
North America and felt it was “‘single
brooded.”’ It overwinters under leaves and
around the tussocks at the edges of swamps
(Torre-Bueno 1925); small colonies have
been found beneath logs along the borders
of woodlands in late October (Blatchley
1926):
90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
This lygaeid has been reared in the lab-
oratory under unspecified conditions and
the egg and nymphal instars, except the sec-
ond, have been described (Slater 1951).
In this paper, we present information on
the field life history in southern Illinois and
laboratory rearing of O. dorsalis and de-
scribe the immature stages.
MATERIALS AND METHODS
Field life history.—During summer
1994, several adults were observed feeding
and copulating on the spikes of Carex crin-
ita Lamarck in the La Rue-Pine Hills Re-
search Natural Area, Union County, IL. The
site is located along the south side of Forest
Road 345, approximately 1.25 miles (2.01
km) east of State Highway 3. The plants
(approximately 80 individuals) were grow-
ing at the bases of red maple (Acer rubrum
L.) located at the edge of La Rue Swamp.
The population of O. dorsalis, although
small, seemed sufficiently large to permit a
life history investigation of the bug. There-
fore, a study was conducted from June 1994
to November 1995. Counts of all nymphs
and adults observed in the field, and notes
on their activity, were taken weekly from
early April to early November, before the
bugs emerged from and after they entered
Overwintering sites, respectively. Speci-
mens were collected by hand picking.
Nymphs large enough to be identified to in-
star, and adults, were released. Others were
preserved in 70% ethanol and taken to the
laboratory for closer examination. Also,
five C. crinita staminate spikes were taken
to the laboratory weekly and inspected for
eggs.
Species of Carex, other than C. crinita,
that served as host plants were noted from
Pine Hills; Oakwood Bottoms, Jackson Co.;
and Lake on the Campus, Southern Illinois
University at Carbondale, Jackson Co. Bugs
were recorded as feeding if their rostra were
inserted in seeds or if many individuals
were on the same plant.
Potential overwintering sites (e.g., leaf
litter, under bark of fallen trees) were ex-
amined periodically during November—
March.
Laboratory rearing.—Ten adults were
collected on 17 May and on 24 May 1995,
brought to the laboratory, and placed in two
oviposition cages (5 males, 5 females/cage).
Each cage consisted of a 1l-quart (approxi-
mately 0.95-liter) Mason jar with a disc of
moistened filter paper on the bottom. A pis-
tillate spike of C. crinita (collected outside
the study site) served as food and was in-
serted in a vial (8.5 cm length, 2.3 cm diam
at base) containing distilled water stoppered
with cotton; the vial was placed upright in
the bottom of the cage. The cage was closed
with a disc of paper toweling and wire
screening secured with the band of the two-
piece Mason jar lid. A strip of paper tow-
eling (approximately 3 XX 20 cm), which
provided additional area for excrement ab-
sorption, was placed inside the jar with one
end over the lip and held in place by the
band.
The cages were examined daily for eggs,
which were removed and placed on moist-
ened filter paper in the bottom of a petri
dish (approximately 9 cm diam., 2 cm
depth) and covered with the lid. A thin lay-
er of petroleum jelly was applied to the in-
side of the rim to help prevent the first in-
stars from escaping. Additionally, the lids
were secured with two strips of masking
tape to insure a tight fit.
Nymphs (including the firsts) were kept
in petri dishes prepared similarly to those
for eggs (e.g., a ring of petroleum jelly); a
section of a pistillate spike served as food.
The nymphs were grouped by molting dates
to accurately determine the stadia.
Food and filter paper in the cages and
jars were changed, and distilled water was
added, approximately every 2—3 days; the
exception to this was that once oviposition
began, spikes in the cages were replaced
daily because they were destroyed during
examination for eggs.
To obtain information on reproductive
behavior and fecundity (fertility was deter-
mined in laboratory rearing), field-collected
VOLUME 99, NUMBER 1
fifth instars were brought to the laboratory
and placed in Mason jars prepared similarly
to oviposition cages. As they reached
adults, they were segregated by sex in two
additional Mason jars. At the end of a min-
imum of two weeks, six pairs were placed
in petri dishes (one male, one female/dish)
prepared similarly to those used for nymphs
but without the ring of petroleum jelly. Be-
havioral observations were made for peri-
ods of 1—2 hours.
All specimens were kept in incubators
maintained at 26+0.5°C and a 16L:8D pho-
toperiod (approximately 2800 lux).
Descriptions of immature stages.—The
description of each stage is based on ten
individuals. Eggs and first through fifth in-
stars were selected from field collected and
laboratory reared individuals; all had been
preserved in 70% ethanol. Only those that
had not swelled in the alcohol were used in
the descriptions. Drawings were made with
the aid of a camera lucida. Measurements,
in mm, were made with an ocular microm-
eter.
Statistics —Averages are expressed as X
+ SE; standard errors of less than 0.005 are
listed as 0.00. Total developmental periods
for males and females were compared with
Student’s t-test; level of significance was
0:05:
RESULTS AND DISCUSSION
Field life history—Oedancala dorsalis
Overwintered as adults that became active
in late April (Figs. 1, 2), began feeding and
copulating on spikes of maturing Carex
crinita, and reproduced shortly thereafter; it
remained on the host into October. Adults
and nymphs also fed on C. blanda Dewey,
C. cephalophora Muhlenberg, C. comosa F
Boott, C. conjuncta FE Boott, C. crus-corvi
Shuttleworth, C. frankii Kunth, C. granu-
laris Muhlenberg, C. squarrosa L., C. tri-
buloides Wahlenberg, C. vulpinoidea Mi-
chaux, and C. vulpinoidea var. ambigua F
Boott (= C. annectens Bicknell) (Table 1).
One adult and two nymphs were observed
on C. lurida Wahlenberg and C. lupulina
9]
Muhlenberg, respectively, but were not ob-
served feeding.
Eggs were found on C. crinita from mid-
May to early September. They usually were
laid singly, but, sometimes, they were de-
posited side by side in clusters of 2—5. Gen-
erally, they were inserted between the
scales of staminate spikes; occasionally,
they were found in pistillate spikes inserted
between the scale and perigynium. Similar
Ovipositional behavior has been observed in
other species of Oedancala (Baranowski
and Slater 1982, 1989).
The first instars were found from early
June to early September (Figs. 1, 2). They
were observed most commonly near the
bases of the staminate spikes with their yel-
lowish orange abdomens protruding from
between the perigynia. Second instars were
found from mid-June to early September,
third instars from mid-June to early Octo-
ber, fourth instars from late June to mid-
October, and fifth instars from early July to
early October. Second instars through adults
were observed most commonly clinging to
the pistillate spikes, feeding on seeds. If
disturbed, nymphs often would fall to the
ground and remain motionless; adults
sometimes would fly a short distance to an-
other plant or to the ground.
The mating position was end-to-end. No
precopulatory behavior was observed in the
field. Copulating adults were noted from
late May to late June and from mid-July to
early August. Other than mating, little in-
teraction between individuals was ob-
served.
Although adults clearly overwinter, no
individuals were located in the field be-
tween November and March. Overwintered
adults were most abundant from late May
to early July and died off soon thereafter.
New adults appeared in late July as evi-
denced by the appearance of callow adults
(N = 6) and rise in number of adults fol-
lowing the first peak of fifth instars. Based
on the peaks of abundance of the adults,
eggs, and the first-fifth instars (Figs. 1, 2),
and the two periods of copulation (see
92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1st INSTAR
PERCENT OF INDIVIDUALS
3rd INSTAR
(N=74)
4th INSTAR
(N=95)
5th INSTAR
(N=139)
| ever e
during 1995 season in Union Co., IL.
above), O. dorsalis apparently was bivol-
tine in southern Illinois.
Laboratory rearing.—Adults fed and
copulated on the pistillate spikes of C. crin-
ita. Of the 697 eggs laid during this study,
90.2% were inserted in the pistillate spikes
between the scales and perigynia and 1.1%
in the filter paper, and 7.8% were laid on
the cotton and 0.9% on the sides of the jar.
Most were laid singly (68.1%) or side by
side in clusters of 2 or 3 (28.0%), less com-
monly in clusters of 4—7 (3.9%).
The incubation period averaged 13.32
days (Table 2). Eggs were pale green when
laid, turning yellowish in 2—3 days. Pink
eyespots were visible in 5 to 6 days. At this
Field cycle of Oedancala dorsalis. Percent in each sample of total individuals of same stage collected
time, the anterior half of the egg began to
redden; the posterior half remained yellow-
ish. The minute egg burster and the paired
dorsal sclerites of the abdomen were visible
in 8 days.
The first instar emerged through a slit in
the cephalic end of the egg. The ground col-
or was yellowish just after emergence; the
sclerotized areas were pink but darkened to
brown within 3—5 hours.
The first, second, third, fourth, and fifth
stadia averaged 6.34, 5.28, 4.52, 4.02, and
5.52 days, respectively. The total develop-
mental period was 39.00 days (Table 2).
There was no significant difference between
total developmental period for males (37.34
VOLUME 99, NUMBER |
Cue
PERCENT OF INDIVIDUALS
ADULT
(N=797)
1st INSTAR
(N=91)
3rd INSTAR
(N=74)
4th INSTAR
(N=95)
5th INSTAR
(N=139)
Fig. 2:
season in Union Co., IL.
+ 0.35 days, N = 125) and females (36.89
+ 0.32 days, N = 122) (Student’s t-test =
—0.9429, df = 245, p = .3467).
Mortality during the nymphal stadia re-
sulted from incomplete ecdysis and unnat-
ural causes (e. g., drowning in water con-
densation inside the dishes).
Precopulatory behavior was observed be-
tween three pairs of males and females and
lasted 5-10 minutes. In two of the encoun-
ters, the first contact was head-to-head with
both individuals touching antennae in a
slow, irregular, up-and-down motion. Sub-
sequently, the male moved around to the
side of the female while antennating her,
continued until he was behind her, and then
mounted her from the side and extended his
genitalia downward until engaged. In one
of the two pairings, the male tapped the fe-
male with his legs during mounting. After
98
AUGUST |SEPTEMBER| OCTOBER
—————_
Field cycle of Oedancala dorsalis. Percent of individuals of each stage per sample during 1995
5 minutes, while still copulating and re-
maining attached, the male dismounted and
rotated to about an 80° angle; shortly there-
after, the pair assumed a 180° end-to-end
position. During copulation, the female
moved about the petri dish, pulling the
male. Copulation lasted approximately an
hour. In the third encounter, the female ini-
tiated copulation. She antennated the male
and crawled on and off his back repeatedly.
The pair began copulating shortly thereaf-
ter. The male cleaned its rostrum and an-
tennae during copulation. This pair re-
mained in copula for approximately 25
minutes.
The six pairs of adults reproduced in the
laboratory, with the females averaging
53.33 = 13.37 eggs (range, 10-108).
The F, adults produced a second gener-
ation. In addition, fifth instars collected in
94 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Southern Illinois food plants of Oedan-
cala dorsalis.
Stages Collection
Host Taxon Collected* Site”
Cyperaceae
Carex blanda Dewey A PH
C. cephalophora Muhlenberg N, A PH
C. comosa F. Boott N,A Ee. PH
C. conjuncta F. Boott A OB
C. crinita Lamarck N, A ILCs 125
C. crus-corvi Shuttleworth A PH
C. frankii Kunth N, A OB, PH
C. granularis Muhlenberg A PH
C. squarrosa L. N,A OB
C. tribuloides Wahlenberg N,A OB
C. vulpinoidea Michaux N, A OB; PH
C. vulpinoidea Michaux var.
ambigua F. Boott (= C.
annectens Bicknell) N, A OB, PH
4N = nymph; A = adult.
> LC = Lake on the Campus, Southern Illinois Uni-
versity; OB = Oakwood Bottoms; PH = La Rue-Pine
Hills/Otter Pond Research Natural Area.
the field in late July and brought to the lab-
oratory reached adults and subsequently re-
produced. Both of these observations sup-
port our statement that O. dorsalis is bi-
voltine in southern Illinois.
Descriptions of immature stages.—Egg
(Fig. 3A—B): Length, 1.41 + 0.01; width,
0.34 + 0.01. Eggs usually laid singly,
sometimes in small clusters of 2—7; each
elongate and whitish to pale green at ovi-
position, turning yellowish red during mat-
uration. Chorion with longitudinal ridges;
3-7 micropylar processes in ring pattern at
cephalic end.
Nymphal instars: The first instar is de-
scribed in detail, but only major changes
from previous instars are described for sub-
sequent instars. Comparative statements
(e.g., more punctate) refer to previous in-
stars. The length is measured from tip of
tylus to tip of abdomen and the width,
across the widest part of the body. Addi-
tional measurements are given in Table 3.
First instar (Fig. 4): Length, 1.59 + 0.02;
width, 0.66 + 0.01. Body generally yellow-
ish orange in live specimens, paler in pre-
served ones. Sclerotized areas of head, tho-
Table 2. Duration (in days) of each immature stage
of Oedancala dorsalis under controlled laboratory
conditions.
Number
Complet-
ing Cumulative
Stage Stadium Range x += SE mean age
Egg? 621. 10416 19.324, 103 fiaee
Nymph
Ist instar 933 3-10 6.34 + .04 19.66
2nd instar 427 2-12 5.28 + .09 24.94
3rd instar 307 1-15 4.52 = 12 29°46
4th instar 263 2-11 4.02 + .07 33.48
Sth instar 247 2-8 5:52 = 104 739300
4697 eggs were laid.
rax, abdomen, and legs dark brown. Head,
thorax, and abdomen lightly setose; legs
and antennae moderately setose. Body elon-
gate, slightly pyriform, greatest width
across abdominal segments 2-3.
Head narrowed in front, lateral margins
tapering before antennae. Tylus exceeding
juga, apex broad. Compound eyes red; pale
line extending from inner margin of each
eye posteromedially, meeting at midline of
vertex, then extending to posterior margin
of head. Antennae four segmented; seg-
ments | to 3 yellowish, segment 4 brown,
segments white at incisures; segment | ex-
tending beyond apex of tylus; segment 4
acute apically; ratio of antennal segment
lengths approximately 9:10:10:18. Rostrum
four segmented, brown, extending onto ab-
dominal segment 1.
Pro- and mesonota mostly sclerotized;
sclerotized areas concolorous with head, in
form of paired plates, plates separated me-
dially. Pronotal plates subquadrate. Meso-
notal plates subrectangular, posterior mar-
gins generally straight, laterally bending
posteriorly, plates approximately 0.4x
length of pronotal plates at midline. Meta-
notum more membranous; paired sclerites
present, each subelliptical, approximately
0.7X length of mesonotal plates at midline.
Thoracic pleura sclerotized, concolorous
with corresponding notal plates; spiracles
present on posterior margins of pro- and
VOLUME 99, NUMBER 1 95
Fig. 3. Scanning electron micrographs of egg of Oedancala dorsalis. A, Egg. B, Micropylar region.
96 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 3. Measurements (mm)* of Oedancala dorsalis instars.”
Nymph
Ist Instar 2nd Instar 3rd Instar 4th Instar Sth Instar
Body length* 1.59 + 0.02 22025003 2166)== O05 3.81 + 0.06 5.26 + 0.08
Antennal segs.
Ist 0.18 + 0.00 0225 0:00 0327-2) 0/01 0.49 + 0.01 O75 = OO
2nd 0.20 + 0.00 0.26 + 0.00 0.34 + 0.00 0.45 + 0.01 0.61 + 0.01
3rd 0.20 + 0.00 0.25 + 0.00 0.34 + 0.00 0.44 + 0.00 0.59 + 0.01
4th 0.37 + 0.00 0.41 + 0.00 0.50 + 0.01 0.61 + 0.01 0.74 = 0.01
Head width
at eyes 0.43 + 0.00 ODS72== 0101 0.73 + 0.01 0.93 + 0.01 1.14 + 0.01
Synthlipsis O32== 10:00 0.41 + 0.01 OD 27-0101 0.64 + 0.01 Oa 0101
Head length‘ 0.30 + 0.01 0.41 + 0.01 0.50 + 0.01 Om + O'02 0.86 + 0.02
Beak segments
Ist 0.18 + 0.00 0.25 + 0.00 0.31 + 0.00 0.40 + 0.01 0.47 + 0.01
2nd 0.23 + 0.00 0.29 + 0.00 0.35 + 0.00 0.44 + 0.01 O52] OO
3rd On7- = 0100 OH23== 10:00 0.29 + 0.00 0.39 + 0.01 0.45 + 0.01
4th 0.24 + 0.00 0.28 + 0.00 0.34 + 0.00 0.40 + 0.00 0.46 + 0.00
Notal lengths‘
Pronotum 0.20 + 0.01 0.29 + 0.01 0.43 + 0.01 0.69 + 0.02 L.025-= {0101
Mesonotum 0.10 + 0.00 Ont 2== O10 0.31 + 0.01 0.59 + 0.02 0.93 + 0.02
Metanotum 0.10 + 0.00 0.11 + 0.00 0.10 + 0.00 0.10 + 0.00 OMI == 0100
Notal widths
Pronotum 0.49 + 0.01 0.69 + 0.01 0.96 + 0.02 1S6)== 0:03 1.81 + 0.02
Mesonotum 053° 0:01 0.74 + 0.02 1.04 + 0.02 iL Sy Sea O08} 216 20102
Metanotum 0.60 + 0.01 0.80 + 0.02 O92 = 0102 1.00 + 0.04 1:27 = O106
Leg lengths
Profemur 0.30 + 0.01 0.40 + 0.00 05670101 0.85 + 0.01 1.29) == O1O0
Protibia 0.31 + 0.01 OMI OlO (ON SI7/ se OO) 0.80 + 0.01 lipyl a= (0)C0)1
Protarsus 0.22 + 0.00 0.25 + 0.00 03370101 0.43 + 0.01 0.57 + 0.01
Mesofemur 0.29 + 0.01 0.38 + 0.01 OS50E2 0/01 OF 25-2010 ONE = (Ov
Mesotibia 0.30 + 0.00 0.39 + 0.00 054220101 OWS == 0101 1O1F = 0:0)
Mesotarsus 0.21 + 0.00 0.25 + 0.00 (32 22 (OMON 0:42 = 0:01 0.55 + 0.01
Metafemur 0.34 + 0.01 0.44 + 0.01 0.61 + 0.02 0.89 + 0.01 27-0101
Metatibia 0.39 + 0.01 0.51 + 0.01 0.68 + 0.01 0.97 + 0.01 1-355== 0102
Metatarsus 0.25 + 0.01 (AAS) BE (COMO O39==70'01 0.49 + 0.01 0.69 + 0.01
Abd. width 0.66 + 0.01 0.90 + 0.02 122 0104 1.64 + 0.03 1-95) 23)0105
ay + SE; values of <0.005 listed as 0.00.
> Based on 10 individuals of each instar.
© Measured at midline.
mesopleura. Thoracic sterna membranous,
concolorous with membranous areas of
nota. Coxae, trochanters, and femora gen-
erally brownish; tibiae brownish, yellowish
apically; tarsi yellowish, last segment
brownish in apical half; profemora slightly
thickened, ventral margin usually with two
small spines in distal half; tarsi two seg-
mented.
Abdomen with paired scent glands me-
dially on intersegmental lines between seg-
ments 4 and 5, and 5 and 6; each pair sur-
rounded by circular, sclerotized plate; scent
glands visible as median orange masses.
Tergum 8 with pair of small, suboval scle-
rites medially; tergum 9 with posterior two
thirds sclerotized; terga 1-6 each with
sparse, transverse row of setae, 2—8 each
VOLUME 99, NUMBER 1 97
Figs. 4-8. Nymphal stages of Oedancala dorsalis. 4, First instar. 5, Second instar. 6, Third instar. 7, Fourth
instar. 8, Fifth instar.
98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
with single lateral seta, those of 5—7 lon-
gest; terga 2—7 each with faint pseudointer-
segmental line visible. Spiracles ventrolat-
erally on segments 2—8. Sterna 5—9 each
with single medial plate, plates of variable
shape; sterna 5 and 6 each with pair of
small sublateral sclerites; sternum 7 with
sublateral sclerite each side on posterior
margin; setae generally more numerous
than those on corresponding terga.
Second instar (Fig. 5): Length, 2.02 +
0.03; width, 0.90 + 0.02. Sclerotized areas
of head and thorax sparsely and irregularly
punctate.
Ratio of antennal segment lengths ap-
proximately 11:13:13:20.
Thorax with notal plates larger, fused
medially, forming single plates; pale, me-
dian fusion line evident from near anterior
margin of pronotum to posterior margin of
metanotum. Pronotal plate covering prono-
tum. Mesonotal plate nearly covering meso-
notum, subrectangular, posterior margin
concave, approximately 0.5 length of pro-
notum at midline. Metanotum with plates
fused; resulting plate subrectangular, cov-
ering approximately half of metanotum, ap-
proximately 0.3 length of mesonotum at
midline. Paired, small, narrow, transverse
sclerites on intersegmental line between
metanotum and abdominal segment |. Pro-
femora thicker, ventral margin dentate in
distal half with sparse row of teeth, distal
two most prominent.
Abdomen, dorsally, sometimes with three
poorly defined, reddish, longitudinal stripes,
one medial and two lateral; segments 5—7
with lateral setae equal in length to those
on remaining segments. Tergum 9 almost
completely sclerotized.
Otherwise, similar to first instar.
Third instar (Fig. 6): Length, 2.66 +
0.05; width, 1.22 + 0.04. Sclerotized area
of head and thorax more punctate.
Ratio of antennal segment lengths ap-
proximately 16:17:17:25.
Mesonotal plate larger, covering meso-
notum, posterior margin concave medially,
approximately 0.8 length of pronotum at
midline; wing pads visible, small, not ex-
tending beyond posterior margin of meta-
notal plate. Metanotal plate larger, covering
more than two thirds of metanotum, poste-
rior margin straight, approximately 0.3 X
length of mesonotal plate at midline. Inter-
segmental sclerites between metanotum and
abdominal segment 1 sometimes hidden by
metanotum. Thoracic spiracles obscured by
sclerotized pleura. Profemora thicker, teeth
on ventral margin more prominent.
Abdomen with three stripes better de-
fined; lateral setae often paired. Sterna 3—9
with medial sclerites, those on sterna 3—4
paired; intersegmental lines of sterna 1—5
occasionally with paired, transverse, linear,
medial sclerites.
Otherwise, similar to second instar.
Kourth instar (Fig. 7): Length, 3-51
0.06; width, 1.64 + 0.03. Punctures more
numerous. Body width usually greatest at
abdominal segments 2-3.
Head with sclerotized areas generally
yellowish to yellowish brown; dorsum with
posterior margin dark brown to black. Pair
of red ocelli sometimes visible, posterior
and mesad to eyes. Ratio of antennal seg-
ment lengths approximately 10:9:9:12. Ros-
trum extending onto metasternum.
Thorax with sclerotized areas generally
yellowish to brown; pleura with continuous,
longitudinal, dark brown to black stripe, ob-
scure to clearly defined. Pronotum usually
with pair of longitudinal dark-brown to
black stripes. Mesonotum with pair of
stripes, concolorous and continuous with
those of pronotum; mesonotum 0.9 length
of pronotum at midline; wing pads longer,
extending to abdominal tergum 2. Metano-
tal plate larger, completely covering meta-
notum, posterior margin arcuate; metano-
tum approximately 0.2 length of meso-
notum at midline; wing pads evident later-
ally, extending to abdominal tergum 2,
almost entirely covered by mesonotal wing
pads. Meso- and metafemora and some-
times profemora more yellowish; profemo-
ra thicker, ventral margin dentate with three
VOLUME 99, NUMBER 1
prominent subequal teeth distally, separated
by smaller teeth.
Abdomen, dorsally, with three, reddish,
longitudinal stripes now clearly defined;
ventrally, with pair of diffuse, sublateral,
reddish, longitudinal stripes. Tergum 8 with
sclerites larger.
Otherwise, similar to third instar.
Fifth instar (Fig. 8). Length, 5.26 + 0.08;
width, 2.16 + 0.02. Punctures more nu-
merous. Body width usually greatest across
mesothorax.
Head with sclerotized areas yellowish.
Ocelli clearly visible. Antennal segment 1
brown on inner margin, remainder of seg-
ment yellowish; segment 2 yellowish; seg-
ment 3 yellow proximally, reddish brown
to brown distally; segment 4 brownish; ra-
tio of antennal segment lengths 5:4:4:5.
Rostrum yellowish, extending onto poste-
rior margin of mesosternum.
Thorax with sclerotized areas yellowish
to yellowish brown; ventrally with pair of
brown markings on meso- and metathora-
ces. Pronotum with stripes reduced, meso-
notum with stripes still well defined; meso-
notum 0.9X length of pronotum at midline;
metanotum 0.1 length of mesonotum at
midline; meso- and metanotal wing pads
longer, extending onto abdominal tergum 3
or 4. Legs yellowish, ventral margin of pro-
femora dark brown; profemora greatly
swollen, ventral margin dentate with four
prominent subequal teeth, separated by
smaller teeth; tarsi apically, subacute dor-
sally.
Abdomen, ventrally, with two well-de-
fined, longitudinal, sublateral red stripes;
longitudinal red stripe present medially,
varying from diffuse to well defined. Ter-
gum 8 with paired sclerites fused to form
plate almost entirely covering dorsal sur-
face.
Otherwise, similar to fourth instar.
Diagnosis.—The five nymphal instars, in
addition to size, can be separated readily by
the relative lengths of the pro- and meso-
nota; sclerotization of the metanotum; rel-
ative lengths of the lateral abdominal setae;
99
presence or absence of dorsal abdominal
stripes; presence or absence, and degree of
development, of wing pads; and presence or
absence, and number and density, of punc-
tures.
There is a progressive increase in the
length of the mesonotum relative to the pro-
notum through all instars. The first instar
can be separated from later instars by the
lack of a single sclerotized plate on the
metanotum, the relatively longer lengths of
the lateral setae on abdominal terga 5—7,
lack of punctures, and lack of dorsal ab-
dominal stripes. The second instar can be
distinguished from later instars by the lack
of distinct mesonotal wing pads; wing pad
development becomes apparent in the third
instar with progressive development in the
subsequent instars. The third-fifth instars
can be distinguished from each other by the
length of the mesonotal wing pads, which
reach the metanotum in the third instar, ab-
dominal tergum 2 in the fourth, and abdom-
inal tergum 3 or 4 in the fifth. Finally, there
is a progressive increase in the number and
density of punctures in the second-fifth in-
stars.
ACKNOWLEDGMENTS
We thank the following faculty and staff
members of Southern Illinois University at
Carbondale: B. M. Burr and W. G. Dyer,
Department of Zoology, for their critical re-
views of the manuscript; S. C. Mueller, Re-
search Photography and Illustration Facili-
ty, for final drafts of the nymphal-stage il-
lustrations; and S. J. Schmitt and R. D. Tin-
dall, Center for Electron Microscopy, for
the SEM photographs of the eggs. We also
thank zoology graduate student C. S. Bundy
for assistance in the field. We are grateful
to the U. S. Forest Service for granting us
permission to collect in the La Rue-Pine
Hills Research Natural Area, Union Co., IL;
and to M. A. Basinger, Department of Plant
Biology, SIUC, and E. L. Shimp, U. S. For-
est Service, for identification of the sedges.
100 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
LITERATURE CITED
Ashlock, P. D. and A. Slater. 1988. Family Lygaeidae,
pp. 167-245. In Henry, T. J. and R. C. Froeschner,
eds., Catalog of the Heteroptera, or True Bugs, of
Canada and the Continental United States. E. J.
Brill, New York. 958 pp.
Baranowski, R. M. and J. A. Slater. 1982. The Pach-
ygronthinae (Hemiptera: Lygaeidae) of Trinidad
with the description of a new species and notes
on other sedge feeding lygaeids. Florida Ento-
mologist 65: 492-506.
. 1989. The utilization of grasses as host plants
by a species of Oedancala (Hemiptera: Lygaei-
dae) with the description of a new species from
Florida and the West Indies. Florida Entomologist
72: 243-251.
Blatchley, W. S. 1926. Heteroptera or True Bugs of
Eastern North America. Nature Publishing Com-
pany, Indianapolis. 1116 pp.
Slater, J. A. 1951. The immature stages of American
Pachygronthinae (Hemiptera: Lygaeidae). Iowa
Academy of Science 58: 553-561.
Slater, J. A. and J. E. O’Donnell. 1995. A Catalogue
of the Lygaeidae of the World (1960-1994). New
York Entomological Society, New York. 410 pp.
Sweet, M. H. 1960. The seed bugs: a contribution to
the feeding habits of the Lygaeidae (Hemiptera:
Heteroptera). Annals of the Entomological Soci-
ety of America 53: 317-321.
Torre-Bueno, J. R. 1925. Some winter bugs. Bulletin
of the Brooklyn Entomological Society 20: 70.
Uhler, P. R. 1877. Report upon the insects collected
by P. R. Uhler during the explorations of 1875,
including monographs of the families Cydnidae
and Saldae [sic], and the Hemiptera collected by
A. S. Packard, Jr., M. D. Bulletin of the United
States Geological and Geographical Survey of the
Territories 3: 355—475, 765-801, plates 27—28.
Van Duzee, E. P. 1888. Mimicry in Hemiptera. Psyche
5: 27-28.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 101-109
TWO NEW GENERA AND THREE NEW SPECIES OF EUPLECTRINI
(HYMENOPTERA: EULOPHIDAE) FROM THE NEW WORLD
G. A. W. WIJESEKARA AND M. E. SCHAUFF
(GAWW) Maryland Center for Systematic Entomology, Department of Entomology,
University of Maryland, College Park, MD 20742, U.S.A.; (MES) Systematic Entomology
Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture % Na-
tional Museum of Natural History, MRC 168, Washington, DC 20560, U.S.A.
Abstract.—Two new genera (Alveoplectrus, type species A. floridanus W. & S. and
Eurycephaloplectrus, type species E. columbianus W. & S.) and three new species (A.
floridanus, A. truncatus, and E. columbianus) of the tribe Euplectrini (Hymenoptera:
Eulophidae) are described. Euplectrus corumbae Ashmead is reclassified Alveoplectrus
corumbae (Ashmead) comb. n. Alveoplectrus floridanus is a parasite of the larvae of
Alarodia slossoniae (Lepidoptera: Limacodidae). Eurycephaloplectrus colombianus 1s pat-
asitic on larvae of Sibine sp. (Lepidoptera: Limacodidae). The two genera are closely
related and share some synapomorphies with the mainly Old World genus Platyplectrus.
Key Words:
Chalcidoidea, Eulophidae, New World, Alveoplectrus, Eurycephaloplectrus,
Alarodia, Sibine, parasites, Limacodidae
The tribe Euplectrini is unique within
Eulophinae being the only cocoon weaving
ectoparasites. Species in Euplectrini are
united by the possession of an elongate hind
tibial spur. This character has been ob-
served to be somewhat variable. For ex-
ample, the euplectrine genus Metaplectrus
Ferriére has a relatively short hind tibial
spur, and it appears very similar to the ge-
nus Euplectrophelinus Girault (J. LaSalle
personal communication), which is not
placed in the Euplectrini. In the absence of
a thorough phylogenetic analysis of all the
included genera, we continue to define Eu-
plectrini based on this tibial spur character.
Members of the tribe are distributed mainly
in the Old World tropics, although species
of Euplectrus Westwood are found world-
wide. The tribe consists of six genera of
which only two, Euplectrus and Platyplec-
trus Ferriére, are known to occur in the
New World (Bouéek 1988). The tribe is
represented in North America by 13 species
of Euplectrus (Burks 1979) and a single
species of Platyplectrus (Schauff and La-
Salle 1993). In our study on Sri Lankan Eu-
plectrini we reported specimens of Euplec-
tromorpha Girault from the New World
(Wijesekara and Schauff 1994). Although
these specimens possess two submedian ca-
rinae on the propodeum, the defining char-
acteristic of Euplectromorpha, further anal-
ysis led us to conclude that these species
are not properly placed in Euplectromor-
pha. In this paper, we erect two new genera
for these species.
Museum abbreviations are as follows:
National Museum of Natural History,
Washington, D.C. (USNM); The Natural
History Museum, London (BMNH). Ter-
minology for surface sculpture follows Har-
ris (1979).
Bouéek (1988) produced a key to the
genera of Euplectrini. His key is modified
102 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
to include our new genera. Awara Boucéek
(1988) is not included in this key as it does
not properly belong in the Euplectrini (Z.
Bouéek and J. LaSalle, in litt.).
KEY TO THE WORLD GENERA OF
EUPLECTRINI
1. Propodeum with single median carina posterior
tosbasalkcupi(hicsglS) ee ane eee ieee 2
— Propodeum with two submedian carinae; me-
dian carina lacking (Figs. 2, 6, 12, 14) ...... 5
2. Scutellum without sublateral groove (as in Fig.
10); pronotum with transverse carina
Be ec R ES CP PRG ais seri oa dd Ruel Lae Euplectrus Westwood
— Scutellum with distinct sublateral groove (Figs.
5, 13, 14); pronotum variable
3. Hind tibia with a single, relatively short, apical
spur; hind basitarsus subequal in length to sec-
onditarsomete se eee eee Metaplectrus Ferriére
— Hind tibia with 2 distinctly elongate apical
spurs; hind basitarsus much longer than second
LATSOMETE 15 BS 5 ay Sac wae ce SSA S Ne pee wenchar ences 4
4. Submedian area of propodeum divided into
more than four areolae; first tergum as long as
half the length of metasoma ... .Aroplectrus Lin
— Propodeum with undivided submedian area;
first tergum of metasoma variable
mags Es) Otte eet gece aap Platyplectrus Ferriere
5. Scutellum without sublateral groove (Fig. 10);
head relatively broad, more than 1.6 wider
than high; petiole distinct, plate-like in dorsal
view (Fig. 12)
Metter te att Oe Eurycephaloplectrus, new genus
— Scutellum with distinct sublateral groove (Fig.
5); head less than 1.5 wider than high; petiole
transverse, not visible dorsally ............ 6
6. Scutellum with sublateral groove broad, con-
tiguous with the posterior margin (Fig. 5); hind
tibia with a single elongate spur (Fig. 7); hind
tibial apex projected beyond point of attach-
ment of basitarsus (Fig. 7); head with distinct
tentorial depression lateral to clypeus (Fig. 4);
postoccipital carina present (Fig. 1); species
predominantly brownish in color .......
badd Net Pt scence iP i ae aie Alveoplectrus, new genus
— Scutellum with sublateral groove narrow, not
contiguous with the posterior margin (Fig. 14);
hind tibia with one or two spurs; hind tibial
apex not extended beyond point of tarsal at-
tachment; head without facial depressions ad-
jacent to clypeus; species including their ap-
pendages yellowish in color...........
Tt A eee oe Euplectromorpha Girault
Alveoplectrus Wijesekara and Schauff,
new genus
(Figs. 1-8, 15, 16)
Type Species: Alveoplectrus floridanus Wi-
jesekara and Schauff.
Diagnosis.—Female with distinct tento-
rial depression on face near lateral margin
of clypeus (Fig. 4); postoccipital carina
present (Fig. 1); forebasitarsus with distinct
strigil (Fig. 3); scutellum with sublateral
groves which continue along the posterior
margin (Fig. 5); mesepimeron with a dis-
tinct pit located closer to the posterior mar-
gin; dorsellum with posteriorly directed
cup-like carina (Fig. 6); propodeum with
two sublateral carinae that diverge posteri-
orly as a narrow triangle originating from
basal cup (Figs. 2, 6); hind leg with single
tibial spur which is as long as first two tar-
someres together; hind tibial margin ex-
tended beyond tarsal attachment point (Fig.
aD:
Description.—Female. Compound eye
with numerous long silvery setae; malar su-
ture present; clypeus distinct but not de-
marcated by supraclypeal suture (Fig. 4);
face convex; postoccipital carina present;
POL 1.5X longer than OOL; frons and ver-
tex uniformly covered with setae; occipital
area flat compared to concave occiput of
Platyplectrus, with scattered setae; mandi-
bles reduced, not meeting medially and
without teeth; first funicular segment and
clava distinctly longer than other segment
(Figs. 15, 16); scape not reaching level of
anterior ocellus; pronotum uniformly cov-
ered with setae, without transverse carina;
mesoscutum similarly covered with setae;
axilla smooth, without setae; sublateral scu-
tellar grooves contiguous along posterior
margin of scutellum; mesepimeron with a
distinct pit close to posterior mesopleural
margin (Fig. 8); dorsellum medially with
carina in form of posteriorly directed cup
(Fig. 6); propodeum with two submedian
carinae (Figs. 2, 6); petiole transverse; me-
tasoma longer than broad, first tergum cov-
ering more than half length of metasoma;
VOLUME 99, NUMBER 1
103
3
Figs. 1-4.
4, A. truncatus, head.
forebasitarsus with distinct strigil (Fig. 3);
hind tibia with a single elongate spur; apical
margin of hind tibia elongate beyond point
of tarsal attachment (Fig. 7); hind basitarsus
longer than following tarsomeres.
Discussion.—This genus can be distin-
guished from Euplectromorpha and Platy-
plectrus by the presence of the anterior ten-
torial depression, extended hind tibial mar-
gin, and cuplike median carina of the dor-
sellum. In both Euplectromorpha and
Platyplectrus the dorsellum does not pos-
sess a cup-like carina, being either smooth
in most species or irregularly carinated (few
Platyplectrus), and their hind tibial margin
does not extend beyond tarsal attachment
point. The clypeal area is smooth with no
demarcations or depressions in Euplectro-
morpha, whereas in Platyplectrus distinct
supraclypeal carinae demarcate the clypeus,
1-3, Alveoplectrus floridanus. 1, Head, posterior view. 2, Dorsellum and propodeum. 3, Strigil.
and there is no distinct depression near the
tentorial pits.
The single hind tibial spur in Alveoplec-
trus is very distinct. Bouéek (1988) consid-
ered the presence of a single tibial spur as
a good generic character to identify Meta-
plectrus but we have observed some vari-
ation among other species. For example,
Euplectromorpha jamburaliyaensis has a
single hind tibial spur, although most Eu-
plectromorpha species have two hind tibial
spurs (Wijesekara and Schauff 1994).
Based on this, we treat this character as
homoplasious within the tribe. The pres-
ence of two submedian carinae on the pro-
podeum has been regarded as autapomorph-
ic for Euplectromorpha but is also charac-
teristic of the two new genera. Euplectro-
morpha differs from Alveoplectrus by
having fine lateral furrows on the scutellum
104 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 5-8.
sepimeron.
which are not posteriorly contiguous, and
lacking postoccipital and pronotal carinae
and posterior pit of the mesepimeron.
In all three species of Alveoplectrus the
occipital carina is very distinctly developed
as in some Platyplectrus, but we consider
this character to be variable, as we also
have examined single specimens of three
additional undescribed species of Alveo-
plectrus from Central and South America,
two of which do not possess the occipital
carina.
The relationship of Alveoplectrus to other
Euplectrini is difficult to assess. Overall,
species of Alveoplectrus appear similar to
those of Platyplectrus, and the posteriorly
contiguous lateral carina on the scutellum,
setaceous head, pronotum and mesonotum,
and distinct strigil suggest a close relation-
ship with that genus. In addition presence
of a distinct pit on the mesepimeron closer
to the posterior margin of the mesopleuron
also suggest a closer relationship to Platy-
Alveoplectrus truncatus. 5, Scutellum. 6, Dorsellum and propodeum. 7, Hindtibial spur. 8, Me-
plectrus. Sharing of similar propodeal ca-
rinae by Alveoplectrus and Euplectromor-
pha is probably homoplasious.
Etymology.—‘‘Alveo” comes from Latin
‘“‘alveus’” meaning cavity, and refers to the
depressions on the face of species in the
genus.
KEY TO THE SPECIES OF ALVEOPLECTRUS
1. Mesepimeron divided by transverse sulcus
(Fig. 8); frons below anterior ocellus without a
transverse carinae o 2. 4). «4 se ee 2
— Mesepimeron not divided by a sulcus; frons
below anterior ocellus with short transverse ca-
LINACS edo Ramet ee corumbae (Ashmead)
2. Antennal clava truncated; distal 3 flagellom-
eres distinct from others being whitish; occip-
ital carinae without a lamina; posterior end of
propodeum lacks alveole . .truncatus, new species
— Antennal clava slightly asymetrical, not dis-
tinctly truncated; all flagellomeres same color;
occipital carinae with a lamina; posterior end
of propodeum with lateral alveoli .......
Pia SR A eae er ecmee me floridanus, new species
VOLUME 99, NUMBER |
Alveoplectrus floridanus
Wijesekara and Schauff,
new species
(Figs. 1-3, 15)
Female.—Length 1.2—2.0 mm. Color:
Head and mesosoma dark honey brown, an-
tenna, legs except hind coxa, petiole, and
metasoma except small area along lateral
margin of first tergum yellow; hind coxa,
petiole, and posterior lateral area of first ter-
gum brownish but lighter than head and me-
sosoma. Head: Width 1.2 height; intero-
cellar distance 2.5 eye width; frons and
vertex evenly covered with short white se-
tae; occipital and postoccipital carinae pres-
ent (Fig. 1); POL:OOL 5:3; antennal clava
slightly asymmetric; single anellus present;
first funicular segment 1.2 longer than re-
maining funicular segments, which are sub-
equal in length (Fig. 15). Mesosoma: Pro-
notum rectangular, collarlike, uniformly se-
tose; mesoscutum similarly covered with se-
tae; scutellum sculpture asperous; scutellum
longer than broad with distinct lateral fur-
rows contiguous with posterior margin; ax-
illa smooth; mesepimeron divided into lower
and upper mesepimeron by transverse sulcus
terminating as a pit close to posterior me-
sopleural margin; propodeum smooth with
two posteriorly diverging submedian carinae
joining posteriorly with two lateral alveoli
(Fig. 2). Metasoma: Longer than broad, pet-
iole transverse, not visible from dorsal side;
first tergum covers more than half metaso-
mal length. Legs and wings: Hind coxa
smooth; hind basitarsus 2 longer than re-
maining tarsomeres; forewing with
postmarginal vein longer than stigmal vein.
Male.—Unknown.
Host.—Larvae of Alarodia slossoniae
(Lepidoptera: Limacodidae).
Distribution._Florida, Monroe Co.
Types.—Holotype 2 on point with data:
Florida, Monroe Co., Crane Key, 16-IV-76.
Coll. D. Simberloff, ex. larvae Alarodia
slossoniae (USNM). Paratypes: same data
as holotype. (2 2 USNM, 1 2 BMNH).
105
Etymology.—The species epithet refers to
the type locality.
Alveoplectrus truncatus
Wijesekara and Schauff,
new species
(Figs. 4-8, 16)
Female.—Length 1.7—2.0 mm. Color:
head and mesosoma dark honey brown to
black; first three funicular segments, coxae,
and metasoma except lateral part of first ter-
gite brownish; scape, pedicel, and legs ex-
cept coxae yellowish; apical part of mid and
hind femora and tibiae honey colored; an-
tennal clava, fourth funicular, dorsal and
ventral surface of first metasomal tergum
whitish. Head: Width 1.2 X height; inter-
ocular distance 2 eye width; occipital and
postoccipital carinae present; POL:OOL 7:
4; first funicular segment 1.5 longer than
second funicular segment; funicular seg-
ments becoming shorter and broader toward
apex of antenna; clava about 1.6 longer
than the preceding segment and distinctly
truncated (Fig. 16). Mesosoma: Pronotum
rectangular and covered with short brown-
ish setae; mesonotum also covered with
similar setae; scutellum and axilla smooth
(Fig. 5); mesepisternum and mesepimeron
smooth, mesepimeron divided into lower
and upper mesepimeron by a transverse sul-
cus, which terminates as a pit close to pos-
terior mesopleural margin (Fig. 8); propo-
deum with diverging submedian carinae en-
closing a coarse raised median area, other-
wise smooth (Fig. 6); posterior margin of
propodeum without lateral carinae forming
posterior alveole. Metasoma: First tergum
extending over half of the metasoma. Legs
and wings: Hind coxa smooth; single elon-
gate hind tibial spur as long as first two
tarsal segments together; hind basitarsus 2 x
longer than second tarsomere; postmarginal
vein longer than stigmal vein.
Male.—Unknown.
Hosts.—Unknown.
Distribution.—Costa Rica.
Types.—Holotype 2 on point with data:
Costa Rica, Puntarenas, Golfo Dulce 3 Km.
106 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
W. Piedras Blancas, 100m. 3-V-1989 coll.
Hanson (BMNH). Paratypes: same data as
holotype (1 2 USNM); Costa Rica, Puntar-
enas, Gulfo Dulce 3 Km. S. Rincon, 10m.
2-III-1989. coll Hanson (2 2) Costa Rica,
Heredia, Chilamate 75m. XII-1989, III-
1990, Hanson and Godoy (1 ¢), (BMNH).
Etymology.—The species epithet refers
to the truncated clava of the antenna.
Alveoplectrus corumbae (Ashmead),
new combination
Euplectrus corumbae Ashmead, 1904: 517.
Holotype ¢: Brazil, Corumba, H. H.
Smith (USNM 60572).
Female diagnosis.—Head, mesosoma,
and metasoma excluding the dorsum of the
first tergum honey brown; legs yellow; an-
tennal clava, third and fourth funicular seg-
ments brownish with a short transverse ca-
rinae just below anterior ocellus; posterior
margin of dorsal axiller surface with dis-
tinct projection over scutoscutellar suture
toward scutellum; basal cup of the propo-
deum enlarged, with a laminated margin;
mesepimeron not divided into two parts by
a sulcus.
Discussion.—Alveoplectrus corumbae
can be easily distinguished from the Alveo-
plectrus species described above by the
presence of distinct projections of the pos-
terior margin of axilla (straight in A. flori-
danus and without any projection in A.
truncatus), reticulate sculpture of the scu-
tellum (smooth and asprous in A. floridanus
and A. truncatus) and lack of a sulcus on
mesepimeron dividing it into two parts (di-
vided into upper and lower mesepimeron by
a sulcus in both A. floridanus and A. trun-
catus (Fig. 8)).
Alveoplectrus corumbae was described
by Ashmead (1904) from a single female
specimen (USNM type no. 60572, exam-
ined). Vidal Sarmiento and DeSantis (1979)
described a male of A. corumbae reared
from Spodoptera frugiperda (J. E. Smith)
(Noctuidae) as “‘allotype’’.
Eurycephaloplectrus
Wijesekara and Schauff,
new genus
(Figs. 9-12)
Type species: Eurycephaloplectrus colom-
bianus Wijesekara and Schauff.
Diagnosis.—Female head distinctly
broader than high; occiput concave; head,
pronotum and mesoscutum uniformly se-
tose; scutellum smooth without lateral fur-
rows and with two anteriorly contiguous
distinct alveoli separating scutellum from
axillae (Fig. 10); propodeum with posteri-
orly diverging submedian carinae which
originate from basal cup separately; median
carina absent (Fig. 12); petiole wider than
long and distinctly visible dorsally, dorsal
surface smooth and expanded like a plate
(Fig. 12); abdomen shorter than mesosoma.
Description.—Female. Head more than
1.6X as broad as high; vertex and occiput
uniformly covered with short setae; eye
densely ciliated; malar sulcus distinct; clyp-
eus not differentiated; occipital area con-
cave; antennal scrobe distinctly marked;
scape not reaching level of anterior ocellus.
Pronotum and mesoscutum uniformly cov-
ered with short setae; axilla and scutellum
smooth; distinct alveoli between scutellum
and axilla; scutellum without lateral furrow;
each axilla with two pairs of short setae
(Fig. 10); mesepimeron with distinct pit
near posterior margin (Fig. 11); dorsellum
smooth; propodeum with two posteriorly
diverging submedian carinae, which reach
posterior margin of propodeum between ab-
domen and coxal foramina, closer to plica,
making broader angle near basal cup when
compared to Alveoplectrus (Fig. 12); peti-
ole broader than long, platelike, and visible
in dorsal view; metasoma shorter than me-
sosoma; strigil of fore basitarsus not dis-
tinct; hind tibia with two elongate spurs.
Discussion.—Eurycephaloplectrus is
closely related to Alveoplectrus. The syna-
pomorphies for this relationship are 1) uni-
formly ciliated head, pronotum, and meso-
notum and 2) basally separated submedian
VOLUME 99, NUMBER 1
Figs. 9-14.
carina on the propodeum. The distinct pit
close to the posterior margin of the mes-
epimeron is similar to the state found in
Platyplectrus and is considered homolo-
gous to the similar structure in Alveoplec-
trus. However this genus differs from Al-
veoplectrus in lack of sublateral scutellar
furrows, absence of a distinct depression on
the face lateral to clypeus; lack of an elon-
9-12. Eurycephaloplectrus colombianus. 9, Head, posterior view. 10, Scutellum. 11, Mes-
epimeron. 12, Dorsellum, propodeum, and petiole. 13, Platyplectrus sp., dorsum. 14, Euplectromorpha sp.,
dorsum.
gate strigil; presence of two pairs of setae
on each axilla; presence of a dorsally ex-
panded petiole; and presence of two hind
tibial spurs. Species of Alveoplectrus pos-
sess distinct depressions lateral to the clyp-
eus, posteriorly contiguous sublateral fur-
rows on the scutellum, distinct strigil, no
setae on axilla, transverse petiole, and a sin-
gle hind tibial spur. Eurycephaloplectrus is
108 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 15-16. Female antenna. 15, Alveoplectrus
floridanus. 16, A. truncatus.
similar to Euplectrus in lacking lateral fur-
rows on the scutellum but differs from it by
having a distinct malar suture and lacking
a median carina on the propodeum.
Etymology.—‘‘Eurys’’ meaning broad
and “‘cephalo”’ meaning head (both Greek)
together refer to distinctly broad head of
this genus, and is combined with the stem
“*plectrus’’.
Eurycephaloplectrus colombianus
Wijesekara and Schauff,
new species
(Figs. 9-12)
Female.—Length: 1.4—1.5 mm. Color:
Head, ventral and lateral mesosoma, hind
coxa, and petiole honey brown; mesosoma
dorsally much darker than head, almost
black; metasoma except anterior dorsal
area, fore and hind coxae and hind femur
light brown; antennae, legs except all coxae
and hind femur and anterior dorsal area of
metasoma yellow. Head: Width 1.6X
height; interocular distance 2.5 eye width;
mouth margin broad; antennal scrobe dis-
tinctly depressed; vertex and occipital area
uniformly covered with setae; eye densely
setose; occipital and postoccipital carina ab-
sent (Fig. 9); POL:OOL 9:7; occiput con-
cave; first funicular segment 1.2 longer
than second funicular segment; funicular
segments 2—4 subequal in length; clava dis-
tinctly longer than funicular segments. Me-
sosoma: Pronotum collarlike and uniformly
setose (Fig. 9); mesoscutum covered with
similar white setae; mesepimeron not divid-
ed (Fig. 11); dorsellum and propodeum
smooth; basal cup distinctly developed; two
submedian carinae originate separately
from basal cup and diverge posteriorly; pro-
podeal margin laterally with few white se-
tae (Fig. 12). Metasoma: Petiole smooth,
broader than long; dorsal surface of petiole
distinctly expanded and plate-like; metaso-
ma distinctly shorter than mesosoma. Legs
and wings: hind tibia with two elongate
spurs; hind basitarsus slightly longer than
other tarsomeres; wings at rest extended be-
yond apex of metasoma; postmarginal vein
longer than stigmal vein.
Male.—Unknown.
Host.—Larvae of Sibine sp. (Lepidop-
tera: Limacodidae).
Distribution.—Colombia, Tibaitata.
Types.—Holotype 2 on point with data:
Colombia, Tibityata, 21-VIII-1974, Coll.
J.A. Jimenez, “‘Acacias vet’’ ex. Sibine lar-
vae (USNM). Paratypes; 3 females with
same data as holotype (2 2 USNM, 1 2
BMNH).
Etymology.—The species epithet refers
to the type locality.
ACKNOWLEDGMENTS
We thank John LaSalle of the Interna-
tional Institute of Entomology, London, for
valuable advice, John Noyes of The Natural
History Museum, London, for loan of ma-
terial for this study and Gregory Zolner-
wich of Texas A&M University and John
Huber of The Canadian National Collecion
for reviewing the manuscript. We appreci-
ate the work of G. Venable on the plates of
SEMs.
LITERATURE CITED
Ashmead, W. H. 1904. Classification of the chalcid-
flies of the superfamily Chalcidoidea with descrip-
a
VOLUME 99, NUMBER 1
tions of new species in the Carnegie Museum, col-
lected in South America by Herbert H. Smith.
Memoirs of the Carnegie Museum, 1: i—xxi +
225-551
Bouéek, Z. 1988. Australasian Chalcidoidea (Hyme-
noptera): A Biosystematic Revision of Genera of
Fourteen Families, with a Reclassification of Spe-
cies. CAB International Institute of Entomology,
Wallingford. 832 pp.
Burks, B. D. 1979. Family Eulophidae, pp. 967—1022.
In Krombein, K. V., P. D. Hurd, Jr., D. R. Smith,
and B. D. Burks, eds. Catalog of Hymenoptera in
America North of Mexico. Vol. 1. Smithsonian
Institution Press, Washington, D.C. 1198 pp.
Harris, R. A. 1979. A glossary of surface sculpturing.
109
California Department of Food and Agriculture.
Occasional Papers in Entomology 28: 31 pp.
Schauff, M. E. and J. LaSalle. 1993. Nomenclatural
notes on genera of North American Eulophidae
(Hymenoptera: Chalcidoidea). Proceedings of the
Entomological Society of Washington 95: 488-
503.
Vidal Sarmiento, J. A. and L. DeSantis. 1979. Nuevas
citas de Himenopteros para la Republica Argen-
tina. II. Revista de la Sociedad Entomologica Ar-
gentina 34: 19-23.
Wijesekara, G. A. W. and M. E. Schauff. 1994. Re-
vision of the tribe Euplectrini of Sri Lanka (Hy-
menoptera: Eulophidae). Oriental Insects 28: 1—
48.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 110-114
A NEW SPECIES OF GALL MIDGE (DIPTERA: CECIDOMYIIDAE) FROM
ECUADOR ASSOCIATED WITH FLOWERS OF CLAVIJA
(THEOPHRASTACEAE)
RAYMOND J. GAGNE, CORNELIA OTT, AND SUSANNE S. RENNER
(RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S.
Department of Agriculture % National Museum of Natural History MRC-168, Washing-
ton, DC 20560, U.S.A.; (CO, SSR) Institut fiir Spezielle Botanik und Botanischer Garten,
Johannes Gutenberg University, D-55099 Mainz, Germany.
Abstract.—A gall midge new to science, Charidiplosis clavijica Gagné, is described
from specimens found standing on male flowers of Clavija weberbaueri (Theophrastaceae)
in eastern Ecuador. The new species is notable for several unique characters involving
the tarsal claws, the male antenna, and the female ovipositor. The new species is compared
to the widespread and only other described congener, Charidiplosis triangularis (Felt), a
species associated with flowers of cacao.
Key Words:
Clavija is a Neotropical plant genus of
50 species (Stahl 1991). The pollination
and breeding system of Clavija species
have not been studied previously in the
field. Two of us (CO, SSR) are investigat-
ing floral morphology in Clavija in an at-
tempt to determine which species are an-
drodioecious, as suggested by Stahl
(1991). The reproductive biology of Clav-
ija weberbaueri Mez. was studied during
two Stays at the Jatun Sacha Biological Re-
serve in eastern Ecuador in September,
1994 and October, 1995. This plant is an
understory shrub of about 0.8-1.5 m
height, frequent along trails at Jatun Sacha.
Flowers are orange, bowl shaped, ca. 1 cm
diameter, and emit a fruity scent, especially
at night. The chemical composition of the
floral scents of C. euerganea Macbr. and
C. repanda Stahl have been investigated
by Knudsen and Stahl (1994). Scents in
these two species are very similar and con-
sist almost exclusively of sesquiterpenes.
In both years when C. weberbaueri was
Neotropical, Charidiplosis, pollination
studied at Jatun Sacha, numerous individ-
uals were in bloom.
One of us (CO) monitored six shrubs
with bisexual flowers and ten male shrubs
from October until December in both
years. The only visitors seen on the male
flowers during several observation periods,
both during the day and at night until 11
p-m., were gall midges of the new species
described here. The flies remained immo-
bile on the flowers for long periods of
time, except at dusk when some individu-
als were observed to fly. Female flowering
shrubs were extremely rare in both years,
and we did not see any visitors on the few
female flowers we found. Whether these
gall midges pollinate the flowers is still un-
clear.
MATERIALS AND METHODS
The new species was collected by one of
us (CO) in September, 1994 and October,
1995, at the Jatun Sacha Biological Re-
serve, Napo Province, Ecuador. The Re-
VOLUME 99, NUMBER |
serve is located 8 km E of Puerto Misa-
hualli on the eastern slope of the Andes at
about 400 m elevation (01°04’S, 77°36'W).
Mean annual precipitation at Jatun Sacha is
3600 mm and the average yearly tempera-
ture is about 24.5°C. Fresh-caught flies
were killed and preserved in 70% ethanol
and later slide mounted for scientific study
using the method outlined in Gagné (1989,
1994). Specimens of the new species have
been deposited in the Museo Ecuatoriano
de Ciencias Naturales in Quito, Ecuador,
and the National Museum of Natural His-
tory, Smithsonian Institution, (USNM) in
Washington, DC, U.S.A. Terminology for
adult morphology follows usage in Mc-
Alpine (1981).
Charidiplosis clavijica Gagné,
new species
(Figs. 1—7)
Adult.—Head (Figs. 1-2): Eye 9-10
facets long at vertex, connate; facets all
closely adjacent. Vertex with narrow, dor-
sal protuberance bearing 2 long setae.
Frons with 3—4 setae per side. Labella
elongate linear in frontal view, more or
less ovoid in lateral view, with about 15
setae laterally and covered with fine, pliant
setulae mesally. Labrum elongate, basally
with several short, sensory setae, covered
beyond base on dorsum with long, fine,
pliant, setulae, and situated between the la-
bellae. Hypopharynx wide, striated dorsal-
ly and with elongate, setulose ligule ven-
trally. Palpus 4 segmented, with basal pal-
piger. Antenna with 12 flagellomeres, ped-
icel enlarged, longer and wider than both
scape and first flagellomere. Male flagel-
lomeres (Fig. 2) gynecoid, cylindrical,
with very short necks, and becoming suc-
cessively narrower from base to apex; cir-
cumfila appressed, as 2 rings encircling the
flagellomeres, interconnected by 2 longi-
tudinal strands. Female flagellomeres as in
male but all of same width.
Thorax: Scutum with 4, single, sparse
longitudinal rows of setae. Anepisternum
without vestiture. Epimeron with 4—6 setae
111
in longitudinal row. Wing length: d, 1.4—
1.6 mm (n = 5); 2, 1.5—1.6 mm (n = 4);
R,; curved apically, joining C posterior to
wing apex; Rs weak, oblique, situated prox-
imal to mid distance between arculus and
R, apex; M;,, evident on distal two-thirds
length; Cu forked; CuP present before Cu
fork. Fifth tarsomeres (Figs. 3, 7) with ac-
ropod partly hidden from view; claws sim-
ple, thin, curving near basal third; empodia
and pulvilli rudimentary.
Male abdomen: First through sixth ter-
gites entire, rectangular, with mostly single,
continuous, posterior row of setae, 1—3 lat-
eral setae on each side near midlength,
sparse setiform scales covering most of
sclerites, and pair of trichoid sensilla on an-
terior margin. Seventh tergite as for sixth
but posterior row of setae interrupted
mesally. Eighth tergite weakly sclerotized,
anterior pair of trichoid sensilla the only
vestiture. Genitalia (Fig. 5—6): cercus nar-
rowing to rounded apex, with several pos-
terior setae; hypoproct slightly longer than
cercus, tapered gradually from base to very
narrow apex, a pair of setae present apical-
ly; aedeagus longer than gonocoxite, ta-
pered evenly from base to narrow apex,
gonocoxite with short, obtuse mesobasal
lobe, cylindrical beyond; gonostylus elon-
gate, cylindrical, barely tapered from base
to apex, setulose near base, asetulose and
ridged beyond.
Female abdomen: First through seventh
tergites as for male first through sixth.
Eighth tergite square, shorter and narrower
than seventh, with scattered setae on pos-
terior half and anterior pair of trichoid sen-
silla. Eighth sternite present, with scattered
setae and anterior pair of trichoid sensilla.
Intersegmental membrane with scattered se-
tae on all surfaces. Ninth tergum (Fig. 4)
dorsally with elongate, curved setae, later-
ally and ventrally with short, scattered se-
tae. Tenth tergum without setae. Cercus
(Fig. 4) elongate-ovoid, completely setu-
lose, with four apical to apicoventral sen-
sory setae; hypoproct elongate, undivided.
Pupa and larva.—Unknown.
112 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-4.
Holotype.—<d, on male fiower of Clavija
weberbaueri, 6-x-1995, Jatun Sacha, Ec-
uador, C. Ott, deposited in the Museo Ecua-
toriano de Ciencias Naturales in Quito
(MECN).
Other material.—10 3, 10 2, same data
as holotype; 2 ¢, 2 2, same data as holo-
type except [X-1994; specimens deposited
in MECN and USNM.
Etymology.—The name clavijica means
“with Clavija.”
Remarks on taxonomy.—The new spe-
cies belongs in Charidiplosis because of its
narrow, elongate male hypoproct in asso-
ciation with the long, tapering aedeagus,
and, in the female, its setose eighth tergite
and sternite and long ovipositor and cerci.
The species will run to Charidiplosis in the
key to Neotropical genera of Cecidomyiidi
in Gagné (1994).
The only other described species of
Charidiplosis clavijica. 1, Male head (lateral; spheroid mass on top of head is a contaminant). 2,
Male scape, pedicel, and base of flagellum (lateral). 3, Apex of tarsus showing partially recessed claws. 4,
Female ninth abdominal segment and cerci (lateral).
Charidiplosis is C. triangularis (Felt), a
widespread species known from North and
South America and reared from mushrooms
and decaying organic matter, including cow
manure. Larvae of the new species will pre-
sumably also be found feeding in decaying
organic matter. Adults of C. triangularis
have been taken repeatedly in cacao flowers
in Costa Rica (Young 1985, as Aphodiplosis
triangularis), although their role on those
flowers is also unknown.
Charidiplosis clavijica differs from C.
triangularis in several derived character
states of the fifth tarsomere, the male an-
tenna, and the setation of the ninth segment
of the ovipositor. The acropods are largely
hidden from view within the covering sides
of the apex of the fifth tarsomeres, the tarsal
claws are elongate and narrow, and the em-
podia and pulvilli are rudimentary (Figs. 3,
7). Because gall midges stand on the apices
VOLUME 99, NUMBER I
Figs. 5-7.
mere and acropod.
of the tarsi, these derived characters may be
adaptations for locomotion or for grasping
a particular kind of surface. The male an-
tenna (Figs. 1—2) is gynecoid and short, the
pedicel is enlarged, and the flagellum be-
comes gradually narrower from base to
apex. While unique in Cecidomyiinae, this
antenna is reminiscent of that of the fast-
flying and swarming Lestremiinae of the
genera Anarete and Conarete and suggests
the new species may have similar habits.
Foreshortened antennae offer less resistance
to the air than the several times longer male
antennae of C. triangularis and most other
male Cecidomyiidi. The setae of the female
ninth tergum (Fig. 4) are unique for their
long length, and the fact that they are
curved evidently allows the ninth segment
to be retracted readily. We suggest that the
long setae function as additional surface
area for female pheromones produced in the
intersegmental region of the ovipositor and
released when the ninth segment is exserted
(Solinas and Isidoro 1991, Isidoro et al.
1992).
Charidiplosis clavijica. 5, Male genitalia (in part; dorsal). 6, Gonostylus (dorsal). 7, Fifth tarso-
ACKNOWLEDGMENTS
We are grateful to Nit Malikul for pre-
paring the microscopic slides and to Keith
M. Harris, Allen L. Norrbom, Natalia J.
Vandenberg, Allen M. Young, and an anon-
ymous reviewer for their comments on
drafts of the manuscript.
LITERATURE CITED
Gagné, R. J. 1989. The Plant-Feeding Gall Midges of
North America. Cornell University Press, Ithaca,
New York. xiii and 355 pp and 4 pls.
1994. The Gall Midges of the Neotropical
Region. Cornell University Press, Ithaca, New
York. xv and 352 pp.
Isidoro, N., I. H. Williams, M. Solinas, and A. Martin.
1992. Mating behaviour and identification of the
female sex pheromone gland in the brassica pod
midge (Dasineura brassicae Winn.: Cecidomyi-
idae, Diptera). Bollettino dell’ Istituto di Entomo-
logica ““G. Grandi” della Universita degli Studi di
Bologna 47: 27-48.
Knudsen, J. T. and B. Stahl. 1994. Floral odours in
the Theophrastaceae. Biochemical Systematics
and Ecology 22: 259-268.
McAlpine, J. EF 1981. 2. Morphology and terminolo-
gy—adults, pp. 9-63. In McAlpine et al., eds.,
114 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Manual of Nearctic Diptera, Vol. 1. Research nas (Diptera, Cecidomyiidae). Redia 74: 441—
Branch, Agriculture Canada, Monograph 27. vi + 466.
674 pp. Young, A. M. 1985. Studies of cecidomyiid midges
Stahl, B. 1991. A revision of Clavija (Theophrasta- (Diptera: Cecidomyiidae) as cocoa pollinators
ceae). Opera Botanica 107: 1-77. (Theobroma cacao L.) in Central America. Pro-
Solinas, M. and N. Isidoro. 1991. Identification of ceedings of the Entomological Society of Wash-
the female sex pheromone gland in the sorghum ington 87: 49-79.
midge, Allocontarinia sorghicola (Coq.) Soli-
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 115-132
FIVE NEW SPECIES AND A NEW RECORD OF COSTA RICAN LEPTONEMA
GUERIN (TRICHOPTERA: HYDROPSY CHIDAE)
FERNANDO MUNOZ-QUESADA
Department of Entomology; University of Minnesota, St. Paul, MN 55108, U.S.A.
Abstract.—Eighteen species of the genus Leptonema (Trichoptera: Hydropsychidae:
Macronematinae) are reported from Costa Rica. In this paper, five additional undescribed
species of Leptonema from Costa Rica are diagnosed, described and illustrated: L. clorito,
L. flintorum, L. huismanae, L. rafita, and L. tapanti. Their distribution records in Costa
Rica are mapped. Also, L. cheesmanae Mosely is illustrated and recorded from Costa
Rica for the first time.
Resumen.—E| género Leptonema (Trichoptera: Hydropsychidae: Macronematinae) pre-
senta dieciocho especies en Costa Rica. En el presente manuscrito se ofrecen las diagnosis,
descripciones e ilustraciones de cinco especies no descritas de Leptonema presentes en
Costa Rica: L. clorito, L. flintorum, L. huismanae, L. rafita, y L. tapanti. Se trazan en
el mapa los registros de distribucién en Costa Rica de estas especies. Ademas, se informa
y se trazan en el mapa los primeros registros de distribuci6n en Costa Rica de L. chees-
manae Mosely, la cual también es ilustrada.
Key Words:
taxonomy
The genus Leptonema Guérin is one of
the best known and most easily recognized
of the Neotropical caddisflies. The adults
are large (1O—40 mm) with light brown to
light green translucent wings. Some species
have black spots or small areas of dark col-
oration on the forewings. In the New
World, the genus is widely distributed from
southern North America through Central
and South America, including the islands of
the Antilles (Flint et al. 1987). Species also
occur in Africa and Madagascar.
The genus was established by Guérin
(1843) for the Brazilian species Leptonema
pallidum. In 1914, Banks reported Lepto-
nema albovirens (Walker) from Costa Rica,
the first record of the genus for Costa Rica.
Leptonema, Trichoptera, caddisfly, new species, Costa Rica, Neotropics,
Mosely (1933) recorded two additional spe-
cies from Costa Rica in his revision of the
genus. Flint, McAlpine, and Ross (1987)
provided an exhaustive taxonomic review
of the world species, and also considered
phylogenetic and biogeographic aspects.
They described 48 new species, four of
them from Costa Rica, and recorded five
additional species from the country. Hol-
zenthal (1988) added six additional species
records. In total, eighteen species of Lep-
tonema have been recorded in Costa Rica,
(Table 1).
In addition to the described species, five
undescribed species of Leptonema were
found in collections made in Costa Rica
from 1986 through 1992 by R. W. Holzen-
116 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. List of Leptonema species and species groups, defined by Flint et al. (1987), recorded in Costa
Rica, with distribution records as published by Flint et al. (1987) and Holzenthal (1988).
SSS
Species
Distribution
ener ee ee ee eee a
Crassum Group
L. crassum Ulmer 1905
L. divaricatum Flint, McAlpine, Ross 1987
Stigmosum Group
L. tapanti, new species
Plicatum Group
. ekisi Flint, McAlpine, Ross 1987
. flintorum, new species
. fortunum Flint, McAlpine, Ross 1987
. hamuli Flint, McAlpine, Ross 1987
. huismanae, new species
rafita, new species
salvini Mosely 1933
. sinuatum Mosely 1933
. turrialbum Flint, McAlpine, Ross 1987
. vitum Flint, McAlpine, Ross 1987
. woldianum Flint, McAlpine, Ross 1987
Teil alll alll alll Spill alll Gall oxlll ool oll oe
Simulans Group
L. asclepium Flint, McAlpine, Ross 1987
L. campanum Flint, McAlpine, Ross 1987
L. simulans simulans Flint, McAlpine, Ross 1987
Pallidum Group
L. albovirens (Walker) 1852
Complexum Group
L. cheesmanae Mosely 1933, new record
L. clorito, new species
L. complexum Mosely 1933
L. forficulum Mosely 1933
L. furciligerum Flint, McAlpine, Ross 1987
L. intermedium Mosely 1933
Mexico; Guatemala; Honduras; Nicaragua; Costa
Rica: Alajuela, Guanacaste, Heredia, Limon;
Panama; Colombia; Venezuela; Brazil; Peru;
Paraguay; Argentina.
Costa Rica: Limén; Colombia; Venezuela; Ecua-
dor.
Costa Rica: Cartago; Panama.
Costa Rica:
Costa Rica:
Costa Rica;
Costa Rica:
Costa Rica:
Costa Rica:
Costa Rica;
Costa Rica;
Costa Rica:
Costa Rica:
Costa Rica;
Alajuela, Cartago; Panama.
Puntarenas.
Panama.
Cartago; Panama.
Alajuela, Guanacaste.
Alajuela, Cartago, San José.
Panama.
Panama; Colombia.
Alajuela, Cartago.
Puntarenas.
Panama.
Costa Rica: Cartago, San José.
Costa Rica; Panama.
Costa Rica: Alajuela, Cartago, Guanacaste, Punta-
renas, San José; Panama.
USA: Texas; Mexico; Belice; Guatemala; Hondu-
ras; Nicaragua; Costa Rica; Panama; Colombia;
Venezuela; Trinidad & Tobago; Granada; St.
Vincent.
Costa Rica: Alajuela, Guanacaste, Limon, San José;
Panama; Colombia.
Costa Rica: Alajuela.
Costa Rica: Alajuela, Cartago, Limon; Panama.
Costa Rica; Panama.
Costa Rica: Puntarenas.
Costa Rica: Alajuela, Cartago, Heredia, San José;
Panama; Colombia; Ecuador.
thal and his colleagues. In the present paper
these species are diagnosed, described, and
illustrated. Also, L. cheesmanae Mosely is
illustrated and recorded from Costa Rica for
the first time. Terminology used for geni-
talic structures follows that presented by
Flint et al. (1987). This paper represents the
results of an ongoing project, sponsored by
the National Science Foundation and the
University of Minnesota Insect Collection,
to catalog and describe the caddisfly fauna
of Costa Rica. Holotypes of the species de-
scribed are deposited in the collections of
the National Museum of Natural History,
VOLUME 99, NUMBER 1
Smithsonian Institution, Washington, DC
(NMNH). Paratypes and other specimens
examined, are deposited in the collections
of the University of Minnesota Insect Col-
lection, St. Paul, Minnesota (UMSP), the
National Museum of Natural History,
Smithsonian Institution, Washington, DC
(NMNH), and the Instituto Nacional de
Biodiversidad, Santo Domingo de Heredia,
Costa Rica, (INBIO). All specimens are
pinned unless otherwise noted.
Leptonema clorito Munoz-Q.,
new species
(Fig. 1, Map 1)
Diagnosis.—This species belongs within
the complexum Group, as defined by Flint
et al. (1987). It is very similar to Leptonema
cheesmanae Mosely, but can be distin-
guished from that species by the shape of
process ‘“‘d” of the phallic apparatus. In L.
clorito, process ‘“‘d’’ consists of only the
apical arm, which is elongate, slender, di-
rected dorsally at base, and curved apically,
as viewed laterally. In L. cheesmanae, pro-
cess “‘d”’ consists of both apical and basal
arms. In lateral view, the apical arm is elon-
gate, curved, and directed posteriorly, the
basal arm is slender, long, and projected an-
teriorly beyond of the apex of process “‘e”’
and the base of process “‘f’’. Finally, in L.
clorito, the lateral lobes of segment X are
triangular, as viewed dorsally; these are
subquadrate in L. cheesmanae.
Description.—Male: Length of forewing
17 mm. Body sclerites pale yellow. Dorsum
of head pale yellow and with short, light
brown setae. Legs with fine, yellowish se-
tae. Wings light green, translucent; fore-
wing covered with fine, short, yellowish se-
tae, with small rounded patch of brownish
setae Over area around anterior angle of me-
dial cell; apical third of forewing slightly
infuscate. Maxillary palpus with fifth seg-
ment about 3/5 length of basal 4 segments
combined. Process of sternum V@ large,
oval. Genitalia (Fig. 1): Segment IX, as
viewed laterally, narrow, elongate, with
V-shaped dorsal keel. Segment X with wart
117
‘“‘a”’ elongate, base narrow, apex bulbous;
wart ““b-/”’ elongate, fingerlike; wart ‘“‘b-2”
short; wart “‘c’’ absent; lateral lobes, as
viewed dorsally, triangular, projecting pos-
teriorly; as viewed laterally, lateral lobes
rounded, bearing short setae on lateral mar-
gin; ventral margin of segment X with
hooklike lobe. Inferior appendage two seg-
mented, basal segment slightly more than 4
times length of apical segment, widest sub-
apically; apical segment with short setae on
inner margin. Phallus with midsection long,
tubular; apical section complex, bearing
two, tiny, sharply pointed, sclerotized phal-
lotremal sclerites behind process ‘“‘a’’, vis-
ible in dorsal view; process “‘a’’, as viewed
laterally, fingerlike, apex truncate, elevated
and arched over process “‘g’’ and phallotre-
mal sclerites; as viewed dorsally, process
“a” tonguelike, apex truncate, lightly scler-
otized, arising dorsomesally; processes “*b-
7” and “*b-2” short, sharply pointed, lightly
sclerotized, arising apically, and directed
anterodorsally; process “‘c’’ long, slender,
arising subapically, apex pointed, directed
anteriorly, and reaching base of process
‘“‘a’’; basal stalk of process “‘d’’, as viewed
laterally, erect, directed dorsally; apical arm
of process *‘d” curved apically, arising dor-
somesally; as viewed dorsally, basal arm of
process ‘“‘d”’ absent; apical arm bifurcated
basally, its projections pointed, projected
posterolaterally; process ‘“‘e-/’’, as viewed
dorsally, spinelike, short, apex rounded,
arising dorsolaterally and projecting poster-
olaterally; process ‘“‘e-2”’, as viewed dor-
sally, bifurcated, arising dorsomesally, its
projections long, slender, slightly serrated,
arched, projecting anteriorly, with pointed
apices reaching base of process ‘‘f’’; pro-
cess “‘f’’ fingerlike, elongate, arising dor-
somesally, apex rounded, reaching base of
apical arm of process ‘“‘d’’; process “‘g” a
ventrolateral lobe, broad, flat, rounded; as
viewed dorsally, emarginated, with mesal
projection, apex rounded and projected pos-
teriorly; process “‘j”’ absent.
Type material.—Holotype: ¢, COSTA
RICA: Alajuela: Cerro Campana, ca. 6 km
118 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
wart "b-1" wart "b-1" wart "b-2"
1H
E apical arm
“o{" process "d"
coe
iE is
Fig. 1. Leptonema clorito, male genitalia. A, Lateral view. B, Segments IX, X, dorsal view. C, Inferior
appendage, posteroventral view. D, Phallus, lateral view. E, Phallus, dorsal view. EK Phallus, ventral view. G,
Maxillary palpus, lateral view. H, Sternum V, ventral view.
VOLUME 99, NUMBER 1
(air) NW Dos Rios, 10.9°N, 85.4°W, el. 640
m, 15—16.iii.1986, Holzenthal and Fasth
(NMNB#).
Etymology.—Dedicated to the memory
of Dr. Clodomiro Picado Twight (1887-
1944), in recognition of his numerous and
outstanding contributions to the biology of
Costa Rica. Dr. Picado was known affec-
tionately as “‘Clorito.”
Leptonema flintorum Munoz-Q.,
new species
(Fig. 2, Map 2)
Diagnosis.—This species is a member of
the plicatum Group, as defined by Flint et
al. (1987). It is very similar to L. huismanae
n. sp., but differs in the shape and size of
process “‘g”’ of the phallic apparatus. In L.
flintorum, process “‘g”’ is narrow, elongate,
distinctly concave dorsally, with a slightly
rounded and slightly serrated apex, barely
reaching the posterior margin of process
‘bh’. In L. huismanae, process “*g”’ is larg-
er, very broad, slightly concave middorsal-
ly, with a broad, rounded, serrated apex,
generally extending beyond the posterior
margin of process ‘“‘b’’. Additionally, the
midsection of the phallus of L. flintorum is
wider than that of L. huismanae. Also, pro-
cess ‘“‘e’’ of the phallus of L. flintorum is
more robust and conspicuous than the same
process in L. huismanae. The pattern of
brownish setae on the forewing of L. huis-
manae is darker than the pattern in L. flin-
torum. Leptonema flintorum has only been
collected in the southern region of Costa
Rica; L. huismanae has been collected in
the central and northern regions of the
country. Finally, L. flintorum and L. huis-
manae can be separated from L. sinuatum
Mosely by the shape of process “‘a’’ and by
the presence or absence of processes “‘e”’
and ‘‘f’’ of the phallus; these latter two pro-
cesses also separate two the new species
from L. hamuli Flint, McAlpine, and Ross.
In L. flintorum and L. huismanae, process
“a” arises dorsally, is broad, membranous,
emarginate apicomesally and without con-
spicuous, dorsomesal, projections; in L. sin-
119
uatum, process “‘a”’ has a pair of conspic-
uous, dorsomesal, curved projections,
which are directed anteriorly, In L. flinto-
rum and L. huismanae, process “‘e’’ is pres-
ent and process “‘f’’ is absent, as viewed
dorsally. In L. hamuli and L. sinuatum, pro-
cess “‘e’’ is absent and process “‘f”’ is pres-
ent.
Description.—Male: Length of forewing
20—22 mm. Body sclerites pale brown. Dor-
sum of head pale brown with short, light
brown setae. Legs with fine, light brown se-
tae. Wings light brown, translucent; fore-
wing covered with fine, short, brown setae,
with darker brown setae along anal veins,
and transverse band of darker brown setae
over cord. Maxillary palpus with fifth seg-
ment more than % length of basal 4 seg-
ments combined. Process of sternum V
large, oval. Genitalia (Fig. 2): Segment IX,
as viewed laterally, narrow, elongate, with
V-shaped dorsal keel. Segment X with wart
‘“‘qa’’ elongate, base narrow, apex bulbous;
warts “‘b-/]” and “‘b-2” elongate, base nar-
row, apex bulbous; wart “‘c’’ absent; lateral
lobes, as viewed dorsally, sharply pointed,
projecting posteriorly; as viewed laterally,
appearing triangular, bearing short setae on
lateral margin. Inferior appendage two seg-
mented, basal segment more than 3 times
length of apical segment; apical segment
with short setae on inner margin. Phallus
long, tubular; midsection bearing process
‘‘e”’ dorsolaterally, slightly narrower than
apical section of phallus (apical section less
than 1.5 times width of midsection); apical
section bearing two, tiny, sharply pointed,
sclerotized phallotremal sclerites behind
process ‘‘a’’, visible in dorsal view; process
‘a’, as viewed dorsally, broad, membra-
nous, arising dorsally, and emarginate api-
comesally; dorsal lobe of process ‘“‘a”’ as
viewed laterally, with small, lightly sclero-
tized point, directed dorsally; process “*b”’
arising apicoventrally, slender, long, reach-
ing the base of process ‘‘e’’, apex pointed;
as viewed laterally, arched dorsally; process
‘‘e”’ spinelike, robust, conspicuous, arises
dorsolaterally, apex pointed, directed anter-
120 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
COSTA RICA
Map 1.
odorsally (in lateral view, height of midsec-
tion of phallus less than 2.5 times length of
process “‘e’’); process “‘g’’ developed into
narrow, elongate, apicolateral lobe, project-
ing posteriorly, as viewed dorsally, distinct-
ly concave dorsally, apex somewhat round-
ed, lightly sclerotized, barely reaching pos-
terior margin of process “‘b’’; as viewed lat-
erally, slightly serrated on dorsal and
ventral margins; ventrally, with U-shaped,
apicomesal emargination; processes ‘“‘c’’,
ud iar Fandoas absent:
Type material—Holotype: 6, COSTA
RICA: Puntarenas: Rio Bellavista, ca. 1.5
@ cheesmanae
* clorito
Arafita
©1989 Theophilus Britt Griwold
Distribution of Leptonema cheesmanae, L. clorito, and L. rafita.
km NW Las Alturas, 8.951°N, 82.846°W,
el. 1400 m, 16—17.111.1991, Holzenthal,
Munoz, Huisman (NMNH). Paratypes:
COSTA RICA: Puntarenas: same data as
holotype except, | d, 4 2 (UMSP); same
except, trib. Rio Bellavista, Las Alturas
(road to quarry) 8.952°N, 82.848°W, el.
1480 m, 19.111.1991, Holzenthal, Munoz,
Huisman, 1 6 (UMSP).
Etymology.—Named in honor of Dr. Ol-
iver S. Flint and his wife, Mrs. Carol Flint,
in recognition to their great labor in the
study of the Neotropical caddisfly fauna
and their help with the author.
VOLUME 99, NUMBER | 121
process "g"
process "b"
process “e" ,
2F
2E
2D poles)
Fig. 2. Leptonema flintorum, male genitalia. A, Lateral view. B, Segments IX, X, dorsal view. C, Inferior
appendage, posteroventral view. D, Phallus, lateral view. E, Phallus, dorsal view. FE Phallus, ventral view. G,
Maxillary palpus, lateral view. H, Sternum V, ventral view.
Leptonema huismanae Munoz-Q.,
new species
(Fig. 3, Map 2)
Diagnosis.—This species belongs within
the plicatum Group of Flint et al. (1987). It
is closest to L. flintorum n. sp., differing
from that species in the shape and size of
process “‘g’’ of the phallic apparatus, as
well as in the size of process “‘e’’, and of
the width of the midsection of the phallus.
Also, these species can be distinguish by
the pattern of brownish setae on the fore-
wing, as discussed in the diagnosis of L.
flintorum. Finally, L. huismanae can be dis-
tinguished from L. sinuatum Mosely by the
shape of process “‘a’’ and by the presence
or absence of processes “‘e”’ and “‘f”’ of the
phallus, these latter two processes also sep-
arate it from L. hamuli Flint, McAlpine, and
Ross, as described in the diagnosis of L.
flintorum.
Description.—Male: Length of forewing
17-20 mm. Body sclerites pale brown. Dor-
sum of head pale brown with short, light
brown setae. Legs with fine, light brown se-
tae. Wings light brown, translucent; fore-
wing covered with fine, short, brown setae,
with small transverse band of brownish se-
tae over basal third and with longer, darker,
transverse band of brownish setae over cord
and margins of medial cell. Maxillary pal-
pus with apical segment more than 2 length
of basal 4 segments combined. Process of
sternum V large, oval. Genitalia (Fig. 3):
Segment IX, as viewed laterally, narrow,
elongate, with V-shaped dorsal keel. Seg-
ment X with wart “‘a’’ elongate, base nar-
row, apex bulbous; warts “*b-/” and “*b-2”
elongate, base narrow, apex bulbous; wart
‘‘c”’ absent; lateral lobes, as viewed dorsal-
ly, rounded, projecting posteriorly; as
viewed laterally, triangular, bearing short
setae on lateral margin. Inferior appendage
two segmented, basal segment slightly
more than 3 times length of apical segment;
apical segment with short setae on inner
margin. Phallus long, tubular; midsection
bearing process “‘e’’ dorsolaterally, distinct-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ly constricted (apical section more than 2
times width of midsection); apical section
distinctly broader, bearing two, tiny, sharply
pointed, sclerotized phallotremal sclerites
behind process “‘a’’, visible in dorsal view;
process ‘‘a’’, as viewed dorsally, broad,
membranous, arising dorsally, and emargin-
ate apicomesally; dorsal lobe of process
‘‘a’’, as viewed laterally, with small, lightly
sclerotized point, directed dorsally; process
‘“‘b” arising apicoventrally, slender, long,
extending beyond the base of process “e”’,
apex pointed; as viewed laterally, arched
dorsally; process “‘e”’ spinelike, short, aris-
ing dorsolaterally, apex pointed, directed
anterodorsally, but in some specimens pro-
cess ‘‘e’’ very short to minute (in lateral
view, height of midsection of phallus more
than 3 times length of process “‘e’’); process
‘*g’’ developed into large, very broad, api-
colateral lobe, projecting posteriorly, as
viewed dorsally, only slightly concave mid-
dorsally, apex broad, rounded, serrated,
lightly sclerotized, normally extending be-
yond posterior margin of process “‘b’’, but
in some specimens barely reaching beyond
posterior margin of process “*b’’; as viewed
laterally, dorsal, apical and ventral margins
with many robust serrations, lightly scler-
otized; as viewed ventrally, with U-shaped,
apicomesal emargination; processes “‘c’’,
“a> “fe and “7° absent:
Type material—Holotype: ¢, COSTA
RICA: Alajuela: Reserva Forestal San Ra-
mon, Rio San Lorencito and tribs.,
10.216°N, 84.607°W, el. 980 m, 6—
10.i11.1991, Holzenthal, Mufioz, Huisman
(NMNH). Paratypes: COSTA RICA: Ala-
juela: same data as holotype except, 13-
16.vi.1986, C.M. and O.S. Flint, Holzen-
thal, 11 ¢6, 2 2 (NMNH); same except, 2—
4.vii.1986, Holzenthal, Heyn, Armitage, 3
3,3 2 (UMSP); same except, 5—9.vii. 1986,
I. and A. Chacoén, 6 3, 2 2 (UMSP); same
except, 2—6.ix.1986, I. and A. Chacon, 1 6,
1 2 (UMSP); same except, 24—27.11.1987,
I. and A. Chacon, 3 6, 1 2 (UMSP); same
except, 30.iii._l.iv.1987, Holzenthal, Hal-
milton, Heyn, 11 ¢ (4 in alcohol), 5 2
VOLUME 99, NUMBER 1 123
Fig. 3. Leptonema huismanae, male genitalia. A, Lateral view. B, Segments IX, X, dorsal view. C, Inferior
appendage, posteroventral view. D, Phallus, lateral view. E, Phallus, dorsal view. F Phallus, ventral view. G,
Maxillary palpus, lateral view. H, Sternum V, ventral view.
124 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
(UMSP); same except, 1—4.v.1990, Holzen-
thal and Blahnik, 5 ¢ (1 in alcohol), 12 °
(UMSP); same Vexcept;) 28=30.vi1:1990;
Holzenthal, Blahnik, Mufioz, 1 d, 1 @ (in
alcohol) (UMSP); same data as holotype
except, 7 d, 10 2 (UMSP); Rio Sarapiqui,
ca. 2. km: SE ~“Cantblanco, 10:299°N-
84.172°W, el. 710 m, 22.111.1986, Holzen-
thal and Fasth, 1 ¢d (UMSP); same except,
611.1987. I and A. Chacon 2 oo 25>
(UMSP); Rio Agrio, ca. 3.5 km NE Bajos
del Toro, 10.243°N, 84.279°W, el. 1290 m,
20.viii.1990, Holzenthal et al., 1 o
(UMSP); Guanacaste: Parque Nacional
Guanacaste, Rio San Josecito, Est. Mengo
[Estaci6n Cacao], 10.922°N, 85.470°W, el.
960 m, 28—29.vii.1987, Holzenthal, Morse,
Clausen, 8 36, 4 2 (UMSP); same except,
Estaci6n Cacao, lado suroeste del Volcan
Cacao, [10°56’N, 85°26'’W], el. 1000—1400
m, ix—xi1.1989, URCG, R. Blanco, C. Cha-
vez, 3 6 (INBIO); same except, vi.1990, II
Curso de Paratax6nomos, 14 6, 12 2 (IN-
BIO); Z.[ona] P.[rotectora] Tenorio, tribs.
Rio San Lorenzo, 6 km NW Tierras Mor-
enas [Tilaran], 10.61°N, 84.98°W, el. 900 m,
17—19.11.1992, Holzenthal, Mufioz, Kjer, 3
a, O72. CUMSEP):
Etymology.—Named in honor of Jolanda
Huisman, in recognition of her great help
with the Trichoptera of Costa Rica Project
and for her friendship.
Leptonema rafita Munoz-Q.,
new species
(Fig. 4, Map 1)
Diagnosis.—This species is also a mem-
ber of plicatum Group, as defined by Flint
et al. (1987). It is most similar to L. ekisi
Flint, McAlpine, and Ross, L. fortunum
Flint, McAlpine, and Ross, L. salvini Mose-
ly, and L. vitum Flint, McAlpine, and Ross,
differing from those species in the shape of
process ‘‘f” of the phallic apparatus. In L.
rafita, as viewed dorsally, process ‘‘f”’ aris-
es dorsomesally and is round; as viewed lat-
erally, it is short, erect, with an apex mod-
erately rounded and directed dorsally; in L.
ekisi, aS viewed dorsally, process ‘‘f”’ is
elongate, tonguelike, with its apex directed
posteriorly; in L. fortunum, as viewed dor-
sally, process ‘“‘f’’ is somewhat elongate,
with a bifid apex directed posteriorly; in L.
salvini, as viewed dorsally, process “‘f”’ is
oval; as viewed laterally, it is short, and
with its apex slightly rounded and directed
dorsally; and in L. vitum, process “‘f”’ is
absent. Also, process “‘g’’ of the phallus is
unilobed in L. salvini, and it is bilobed in
L. rafita, L. ekisi, L. fortunum and L. vitum.
In addition, process “‘g’’ is different among
the four species. In L. rafita, process “‘g”
is short, lightly sclerotized, arising apico-
laterally, directed posteriorly, and bilobed;
as viewed laterally, the apical lobe of pro-
cess “‘g”’ is erect, subtriangular, strongly
serrated, projected posteriorly, with a pair
of apical points that reach the posterior
margin of the process “‘b’’; the dorsal lobe
of the process “‘g’’ is erect, subtriangular,
slightly serrated, directed dorsally and with
a pointed apex; in L. ekisi, process “‘g”’ is
short, apicolateral, directed posteriorly and
bilobed apically; as viewed laterally, the
apical lobe is short, rounded, unserrated,
projected posteriorly, and barely reaching
the posterior margin of process ‘“*b’’; the
dorsal lobe is subtriangular, directed dor-
sally with small apical points; in L. fortun-
um, process “‘g’’ is elongate, apicolateral,
projected posteriorly beyond the posterior
margin of process “‘b’’, and bilobed apical-
ly; as viewed laterally, the apical lobe is
subtriangular, unserrated, with a pointed
apex and directed posteriorly; the dorsal
lobe is erect, subtriangular, unserrated, with
a pointed apex and directed dorsally; in L.
salvini, process “‘g”’ is unilobed, short, ap-
icolateral; as viewed laterally, projected
posteriorly, reaching the posterior margin of
process “*b’’; the apical lobe is absent; and
the dorsal lobe is erect, subtriangular, dor-
soapical, and directed dorsally; and in L.
vitum, process “‘g”’ is elongate, apicolateral,
projected posteriorly beyond of posterior
margin of process “‘b’’, and bilobed; the
apical lobe, as viewed laterally, is large,
quadrate, serrated posterodorsally and di-
VOLUME 99, NUMBER |
rected posteriorly; as viewed dorsally, with
apex directed posteromesally and the dorsal
emerging mesally; the dorsal lobe, as
viewed laterally, is erect, slender, and di-
rected dorsally. Finally, in both L. rafita and
L. salvini, process ‘“‘e’’ of the phallus, as
viewed laterally, reaches the base of the
process “‘f’’; in L. ekisi, it does not reach
the base of process “‘f”’ and in L. fortunum,
it is absent. Finally, in L. vitum. process
“7 1S absent.
Description.—Male: Length of forewing
20—23 mm. Body sclerites pale brown. Dor-
sum of head pale brown with short, light
brown setae. Leg segments with fine, light
brown setae. Wings light brown, translu-
cent; forewing covered with fine, short,
brown setae, with small, rounded patch of
brownish setae over posterior margin of
medial cell. Maxillary palpus with apical
segment more than 2 length of basal 4 seg-
ments combined. Process of sternum V
large, oval. Genitalia (Fig. 4): Segment IX,
as viewed laterally, narrow, elongate, with
V-shaped dorsal keel. Segment X with wart
‘qa’? short with short setae; wart “b-/”
short with short setae; wart ““b-2” elongate,
base narrow, apex bulbous; wart “‘c”’ ab-
sent; lateral lobes, as viewed dorsally,
rounded, projecting posteriorly, bearing
short setae; as viewed laterally, rounded,
bearing short setae on lateral margin. Infe-
rior appendage two segmented, basal seg-
ment more than 3 times length of apical
segment; apical segment with short setae on
inner margin. Phallus with midsection long,
tubular; apical section bearing two, tiny,
sharply pointed, sclerotized phallotremal
sclerites behind process ‘‘a’’, visible in dor-
sal view; process “‘a’’, as viewed dorsally,
subtriangular, membranous, arising dorso-
mesally, and emarginate apicomesally; as
viewed laterally, with margin rounded; pro-
cess “b” long, broad, tubular, sinuous,
emerging apicoventrally, apex rounded,
with small point, reaching base of process
“f’, posteroventral section serrated, lightly
sclerotized, as viewed laterally, arched, el-
evated over process “‘e’’, projecting anteri-
orly; process ‘‘e” dorsolateral, fused to
phallobase, with elongate, lateral row of
spicules and ending in membranous, round-
ed, dorsolateral, spiculose lobe, reaching
base of process “‘f’’, as viewed laterally;
process ‘‘f’’, as viewed dorsally, rounded
and arising dorsomesally; as viewed later-
ally, short, erect, apex moderately rounded,
directed dorsally; process “‘g’’ apicolateral,
short, lightly sclerotized, directed posteri-
orly, bilobed; apical lobe of process *“‘g”’ as
viewed laterally, subtriangular, erect,
strongly serrated, projected posteriorly,
with pair of apical points reaching posterior
margin of process ‘‘b’’; dorsal lobe of pro-
cess ‘‘g’’ subtriangular, erect, slightly ser-
rated, with apex pointed directed dorsally;
ventrally, with deep, serrated, U-shaped,
apical emargination; processes “‘c”’, “‘d”
and ‘‘j’’ absent.
Type material—Holotype: ¢, COSTA
RICA: Alajuela: Rio Peje and falls, ca.
1km SE San Vicente, Ciudad Quesada,
10.277°N, 84.388°W, el. 1450 m, 14-
15.11.1992, Holzenthal, Mufioz, Kjer
(NMNH). Paratypes: COSTA RICA: Ala-
juela: Rio Toro, 3.0 km (road) SW Bajos
del Toro, 10.204°N, 84.316°W, el. 1530 m,
11.11.1992, Holzenthal, Mufioz, Kjer, 1 d
(UMSP); same data as holotype except, 4
3,4 2 (UMSP); Cartago: Reserva Tapanti,
Rio Grande de Orosi, 9.686°N, 83.756°W,
el. 1650 m, 15—16.vii.1987, Holzenthal,
Morse, Clausen, | d (UMSP); same except,
Quebrada Palmitos and falls, 9.72°N,
83.78°W, 1400 m, 1-2.vili.1990, Holzen-
thal, Blahnik, Mufioz, 1 ¢, 1 2 (UMSP);
same except, 21.11.1992, Holzenthal, Mu-
noz, Kjer 166, 1 2 (NBIO); sameexcept,
waterfalls, ca. | km (road) NW tunnel,
9.69°N, 83.76°W, 1600 m, 2-3.viii.1990,
Holzenthal, Blahnik, Mufioz, 1 d (NMNH);
San José: trib. to Quebrada Caraigres 3.6
kim! @oad) SW «La Wesuay 9:728°N,
S41 25°W, el 1650 im; 23221992, Holzen-
thal, Kjer, Quesada, 2 6 (UMSP).
Etymology.—Named in honor of Dr.
Ralph W. Holzenthal, who first introduced
me to the Neotropical caddisfly world.
126 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
process "f"
Fig. 4. Leptonema rafita, male genitalia. A, Lateral view. B, Segments IX, X, dorsal view. C, Inferior
appendage, posteroventral view. D, Phallus, lateral view. E, Phallus, dorsal view. F Phallus, ventral view. G,
Maxillary palpus, lateral view. H, Sternum V, ventral view.
VOLUME 99, NUMBER 1
COSTA RICA
Lambert Conformal Come Projection
Map 2.
Also, Dr. Holzenthal has added enormously
to the knowledge of the caddisfly fauna of
Costa Rica, where he is called affectionate-
ly ‘“‘Rafita.”’
Leptonema tapanti Munoz-Q.,
new species
(Fig. 5, Map 2)
Diagnosis.—Leptonema tapanti is the
first species of the stigmosum Group of
Flint et al. (1987) recorded from Costa
Rica. This new species is close to L. auri-
culatum Flint, McAlpine, and Ross of the
same group, but is also similar to L. mich-
A flintorum
@ huismanae
* tapanti
€1989 Theophilus Brit Grepwrold
Distribution of Leptonema flintorum, L. huismanae, and L. tapanti.
ocanense Flint, McAlpine, and Ross, L. pli-
catum Mosely, and L. simplex Mosely of
the plicatum Group. However, it can be dis-
tinguished from those species by the shape
of processes ‘‘b’”’ and “‘c” of the phallus.
In L. tapanti, processes “‘b” and ‘‘c”’ are
fused into a lightly sclerotized, apicolateral,
inflated, paired lobes; as viewed dorsally,
process “‘b-c’’ is earlike, enlarged, and con-
cave dorsolaterally at its middle. The lateral
margin is convexly curved, with the apices
pointed and directed posteriorly, and with a
slender, curved, midlateral projection. In L.
auriculatum, L. michoacanense, L. plica-
128 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tum, and L. simplex, processes “‘b”’ and “‘c”’
are not fused. In L. auriculatum, process
‘*‘b’’ is a small dorsal angle; process “‘c”’ is
smaller, rounded, and earlike; in L. michoa-
canense, process “‘b” is long, slender, ba-
sally curved and with a pointed apex; pro-
cess “‘c’”’ is shorter, slender, curved basally;
in L. plicatum, process “‘b”’ is long, slender,
curved basally and with a pointed apex;
process ‘“‘c’’ is absent; and in L. simplex,
process “*b”’ is short; as viewed laterally, it
is subtriangular with a pointed apex direct-
ed anterodorsally; process “‘c’’ is absent. Fi-
nally, in this new species, as viewed dor-
sally, process “‘b-c”’ has a slender, curved,
midlateral projection, with the apex pointed
and directed mesally; a similar projection is
absent in the other species.
Description.—Male: Length of forewing
17-20 mm. Body sclerites pale brown. Dor-
sum of head pale brown with short, light
brown setae. Leg segments with fine, light
brown setae. Wings light brown, translu-
cent; forewing sparsely covered, mainly be-
tween anal veins, with fine, short, brown
setae, with two small rounded patches of
darker setae, one over each nygma. Maxil-
lary palpus with fifth segment about %
length of basal 4 segments combined. Pro-
cess of sternum V very small, circular. Gen-
italia (Fig. 5): Segment IX, as viewed lat-
erally, narrow, elongate; as viewed dorsally,
with V-shaped dorsal keel, dorsal margin
slightly projecting posteromesally. Segment
X with wart ‘“‘a” absent; warts “‘b-/” and
‘“*b-2”’ short with small setae; wart ‘“‘c”’ ab-
sent; lateral lobes, as viewed dorsally, each
with pair of apical points, projecting pos-
teriorly; as viewed laterally, bearing short
setae on lateral margin. Inferior appendage
two segmented, basal segment less than 3
times length of apical segment; apical seg-
ment with short setae on inner margin.
Phallus with midsection long, tubular; api-
cal section bearing two, tiny, sharply point-
ed, sclerotized phallotremal sclerites behind
process “‘a’’, visible in dorsal view; process
“‘a’’, as viewed dorsally, broad, rounded,
membranous, arising dorsomesally, and
emarginate apicomesally, trilobed; lateral
lobes of process ‘“‘a’’ broad, with lateral
margin slightly convex; mesal lobe of pro-
cess ‘“‘a’’ slender; as viewed laterally, lat-
eral lobes of process “‘a’’, rounded and di-
rected dorsally; mesal lobe of process “‘a”’
rounded, higher, directed anterodorsally,
and reaching posteromesal spicules of mes-
al row of spicules of process ‘“‘e’’; processes
‘“*b” and ‘‘c”’ fused into lightly sclerotized,
apicolateral, inflated, paired lobes; as
viewed dorsally, process “‘b-c”’ earlike, en-
larged, concave dorsolaterally at middle,
lateral margin convexly curved, apices
pointed, directed posteriorly, and with slen-
der, curved, midlateral projection with
pointed apex, directed mesally; ventrally,
with deep, narrowly V-shaped, apicomesal
emargination; process ‘‘e’’ dorsolateral,
fused to phallobase; as viewed dorsally, de-
veloped into three elongated rows of spic-
ules, one mesal and Y-shaped, and two lat-
eral ones ending in rounded, membranous,
dorsal, and anteriorly directed lobes, with
small spicules; processes “adi ff same
and *‘7’’ absent.
Type material.—Holotype: ¢ COSTA
RICA: Cartago: Reserva Tapanti, Quebra-
da Palmitos and falls, 9.72°N, 83.78°W,
1400 m, 24—25.i11.1991, Holzenthal, Mu-
fioz, Huisman (NMNH). Paratypes: COSTA
RICA: Cartago: same data as holotype ex-
cept, Rfo Grande de Orosi, 9.686°N,
83.756°W, el. 1650 m, 18—21.i11.1987, Hol-
zenthal, Hamilton, Heyn; i Goes
(UMSP); same except, Rio Dos Amigos
and falls, ca. 6 km (road) NW tunnel,
9.704°N, 83.783°W, 1500 m, 23.11i.1991,
Holzenthal, Mufioz, Huisman 1 6, 1 &
(UMSP); same data as holotype except, 10
3, 2-2 CUMSP); same exeept, Zien 19928
Holzenthal, Mufioz Kjer, 1 ¢d, 3 2 (INBIO).
PANAMA: Chiriqut: Guadalupe Arriba,
8°52'26"N, 82°33'13"W, 1—28.11.1984, H.
Wolda, 3 6, 4 2 (NMNH); same except,
29.ii.—27.ii1.1984, 3 d, 5 2 (NMNH); same
except, 3—30.iv.1984, 1 ¢6, 8 2 (NMNH);
same except, 2—29.v.1984, 2 6, 3 2
(NMNH); same except, 30.v.—19.vi.1984, 2
VOLUME 99, NUMBER 1 129
SG
process "b-c"
5H
process "e"
Fig. 5. Leptonema tapanti, male genitalia. A, Lateral view. B, Segments IX, X, dorsal view. C, Inferior
appendage, posteroventral view. D, Phallus, lateral view. E, Phallus, dorsal view. F Phallus, ventral view. G,
Maxillary palpus, lateral view. H, Sternum V, ventral view.
130
6 (NMNH); same except, 15—28.v.1985, 2
3d 2, 2 \(NMNBE); ‘same excepinis—
18.vi.1985, 3 d6 (NMNH); same except,
14—29.viii.1985, 1 56, 1 2 (NMNH).
Etymology.—Named for the type locali-
ty, Tapanti National Park, which has a rich
and endemic caddisfly fauna.
Leptonema cheesmanae Mosely
(Fig. 6, Map 1)
Leptonema cheesmanae Mosely 1933: 51-—
52, figs. 148-153, 3, I. Gorgona, Colom-
bia (BMNH); McElravy et al. 1981: 153;
19827307"
Distribution.—Colombia, Panama, Costa
Rica (new record).
Diagnosis.—This species is a member of
the complexum Group, as defined by Flint
et al. (1987). Leptonema cheesmanae is
very similar to L. clorito n. sp., differing
from that species in the shape of process
‘“‘d”’ of the phallic apparatus and the shape
of the lateral lobes of segment X, as viewed
dorsally, as described in the diagnosis of L.
clorito. Also, this species can be separated
from L. harpagum Flint, McAlpine, and
Ross by the shape of process ‘“‘a’’, which
in L. cheesmanae, is unilobed, as viewed
dorsally, process “a” is tonguelike, elon-
gate, slender and arising dorsomesally; as
viewed laterally, it is fingerlike, arched,
with the apex truncated. In L. harpagum,
process ‘“‘a” is bilobed and prominent. In
figure 153 from Mosely (1933) of L. chees-
manae, process “‘f”’ is not illustrated, but it
does appear in his figure 152. However, in
figure 152 it is not possible to see process
*“‘e-1’’, only process ‘“‘e-2”’.
New distribution records.—COSTA
RICA: Alajuela: Cerro Campana, ca. 6 km
(air) NW Dos Rios, 10.9°N, 85.4°W, el.
640 m, 15—16.111.1986, Holzenthal and
Fasth, 1 d6 (UMSP); Reserva Forestal San
Ramon, Rio San Lorencito and tribs.,
10.216°N, 84.607°W, el. 980 m, 2-
4.vi1.1986, Holzenthal, Heyn, Armitage, 5
3 (UMSP); same except, 5—9.vii.1986, I.
and A. Chacon, 1 ¢ (UMSP); same except,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
2—6.ix.1986, I. and A. Chac6én, 9 ¢, 11 2
(UMSP); same except, 1—4.vii.1986, I. and
A. Chac6ny52"ds i 2 GU MSP); ‘same tex-
cept, 24—27.11.1987, I. and A. Chacon, 2 3
(UMSP); same except, 30.iii1.1987, Hol-
zenthal, Hamilton, Heyn, 2 6 (in alcohol)
(INBIO); same except, 1—4.v.1990, Hol-
zenthal, Blahnik, 28 d (10 in alcohol), 3
2 (UMSP); same except, 28—30.vii.1990,
Holzenthal, Blahnik, Mufioz, 3 6, 12 @
(UMSP); same except, 6—10.i11.1991, Hol-
zenthal, Mufoz, Huisman, 4 6, 9 9
(UMSP); Guanacaste: Parque Nacional
Guanacaste, Estacion Pitilla, Rio Orosi,
10.991°N, 85.428°W, el. 700 m, 22-
25.v.1990, Holzenthal and Blahnik, 1 6
(UMSP); same except, Estacién Maritza,
Rio Tempisquito, 10.958°N, 85.497°W, el.
550 m, 16.11.1994, E Mufioz-Q., 4 6,3 2
(in alcohol) (INBIO); Limon: Reserva
Bioldgica Hitoy Cerere, Estaci6n Miramar,
9.671°N, 83.030°W, el. 550 m, 11.xii.1990,
E Mujfioz-Q., 1 ¢ (UMSP); San José:
P.[arque] N.[acional] Braulio Carrillo,
Est.[aci6n] Carrillo, Q.[uebrada] Sangui-
juela, 10.160°N, 83.963°W, el. 800 m, 22—
28.viii.1986, I. and A. Chacon, 3 6,1 @
(UMSP); same except, 27.11i1.1987, Hol-
zenthal, Hamilton, Heyn, 4 6 (UMSP).
ACKNOWLEDGMENTS
Special thanks to Dr. Oliver S. Flint, Jr.,
Dr. Ralph W. Holzenthal, and Dr. Roger J.
Blahnik, who reviewed the manuscript and
made many useful suggestions. Also, I
would like to express thanks to the Gradu-
ate School, University of Minnesota and the
Consejo Nacional para Investigaciones
Cientificas y Tecnol6gicas (CONICIT), San
José, Costa Rica for the support given. This
material is based upon work supported by
National Science Foundation grants BSR-
8917684 and DEB-9400632. This is paper
number 22,375 of the Scientific Journal Se-
ries, Minnesota Agricultural Experiment
Station, St. Paul, Minnesota.
VOLUME 99, NUMBER 1 131
lp
6G
process "d"
basal arm
6E 6F
process "e-2"
6D
Fig. 6. Leptonema cheesmanae, male genitalia. A, Lateral view. B, Segments IX, X, dorsal view. C, Inferior
appendage, posteroventral view. D, Phallus, lateral view. E, Phallus, dorsal view. E Phallus, ventral view. G,
Maxillary palpus, lateral view. H, Sternum V, ventral view.
132 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
LITERATURE CITED
Banks, N. 1914. Neuroptera and Trichoptera from
Costa Rica. Entomological News 25: 149-150.
Flint, O. S., Jr, J. E McAlpine, and H. H. Ross. 1987.
A Revision of the Genus Leptonema (Trichoptera:
Hydropsychidae: Macronematinae). Smithsonian
Contributions to Zoology 450: 1-193.
Guérin-Meneville, M. FE E. 1843. Trichoptera. Jn
Iconographie du Régne Animal de Cuvier 3b:
395-397.
Holzenthal, R. W. 1988. Catalogo Sistematico de los
Tric6pteros de Costa Rica (Insecta: Trichoptera).
Brenesia 29: 51-82.
McElravy, E. P., V. H. Resh, H. Wolda, and O. S. Flint,
Jr. 1981. Diversity of adult Trichoptera in a “‘non-
seasonal” tropical environmental., pages 149—
156. In Moretti, G. P., ed., Proceedings of the 3rd
International Symposium on Trichoptera. Junk,
The Hague.
McElravy, E. P., H. Wolda, and V. H. Resh. 1982.
Seasonality and annual variability of caddisfly
adults (Trichoptera) in a ““Non-Seasonal” Tropical
Environmental. Archiv fiir Hydrobiologie 94:
302-317.
Mosely, M. E. 1933. A Revision of the Genus Lep-
tonema. British Museum (Natural History), Lon-
don, 1-69.
Ulmer, G. 1905. Zur Kenntniss aussereuropdischer
Trichopteren. Stettiner Entomologische Zietung
66: 1-119.
Walker, E 1852. Catalogue of the Neuropterous In-
sects in the Collection of the British Museum, Part
I: Phryganides - Perlides. London. 192 pages.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 133-135
A NEW AMMOPLANUS GUSSAKOVSKIJ
(HYMENOPTERA: SPHECIDAE: PEMPHREDONINAE) FROM
SAN CLEMENTE ISLAND, CALIFORNIA
ARNOLD S. MENKE
Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart-
ment of Agriculture, % National Museum of Natural History, MRC 168, Washington, DC
20560, U.S.A. Current address: Ammophila Research Institute, 1429 Franklin Street, Bis-
bee, AZ 85603, U.S.A.
Abstract—Ammoplanus clemente, n. sp., is described from a single female from San
Clemente Island, California. This is the first record of the genus from the Channel Islands
of California.
Key Words:
In North America, the genus Ammoplan-
us Gussakovskij as defined by Bohart and
Menke (1976) is represented by seven de-
scribed species. Except for A. unami Pate
from Pennsylvania, Virginia, and West Vir-
ginia, the species occur in the western Unit-
ed States, especially in southern California.
Pate (1943) provided an illustrated key to
them, but little has been done with the ge-
nus in North America since his work.
Krombein (1956) described the formerly
unknown male of A. unami. Only two spe-
cies are known from both sexes, A. che-
mehuevi Pate and A. unami. Ammoplanus
loti Pate, A. quabajai Pate, and A. vanyumi
Pate are known by females, and A. sechi
Pate and A. tetli Pate by males.
Marshakov (1979) treated the genus Am-
moplanellus Gussakovskij sensu Bohart and
Menke (1976) as a subgenus of Ammoplan-
us. Marshakov’s action probably has merit
but I have not investigated the matter. Am-
moplanellus differs from Ammoplanus in
having the marginal cell open along the
wing margin, and thus none of the four
North American species of Ammoplanellus
sensu Marshakov can be confused with the
new species of described below.
Ammoplanus, clemente, San Clemente Island, California Channel Islands
Ammoplanus is unrecorded from the Cal-
ifornia Channel Islands (Rust et al. 1985).
The purpose of this paper is to describe a
new species from San Clemente Island so a
name will be available to workers studying
this insular fauna.
The holotype is deposited in the National
Museum of Natural History, Washington,
D.C. Sculpture terminology used here is
from Harris (1979).
I thank Scott Miller, Bishop Museum,
Honolulu; Alex Antropov, Moscow State
University, Moscow; R. M. Bohart, Uni-
versity of California, Davis; W. J. Pulawski,
California Academy of Sciences, San Fran-
cisco; and Steve Nakahara and Ron Hodg-
es, Systematic Entomology Laboratory,
USDA, Beltsville, MD, and Washington,
DC, for reviewing the manuscript.
Ammoplanus clemente Menke,
new species
Holotype female.—Black; mandible pale
amber, scape amber, clypeal lobe reddish
amber, tarsi brown; stigma uniformly, costa
and subcosta of forewing dark brown, other
veins paler. Body sparsely setose except se-
134 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-2. Ammoplanus clemente, holotype; un-
coated SEM photographs. 1, Face. 2, Clypeal notch.
tae denser on legs and apex of gaster. Facial
proportions and shape as in Fig. 1; clypeus
with broad, parallel-sided, U-shaped emar-
gination at base of which a tiny tubercle
bears a long seta that projects into emargi-
nation (Figs. 1—2); emargination bounded
distally by pair of acuminate lobes (Figs. 1—
2); clypeal surface polished, smooth; lower
frons coriaceous, upper frons, vertex and
gena smooth, polished, sparsely, shallowly
punctate (punctures 2 or more diameters
apart, mostly 4 or more); midline of frons
deeply impressed at level of eye tops (does
not show in Fig. 1); frons along inner orbit
at eye top with deep, narrow impression
that delimits an oval area next to eye (Fig.
1); labrum longer than wide, with narrow
U-shaped emargination; mandible apically
bidentate; ventral surface of head shallowly,
transversely concave at level of hypostomal
apex; gena between mandible socket and
occipital carina vaguely microstrigulate.
Pronotum coriaceous; scutum, scutellum,
metanotum, and mesopleuron polished al-
though scutum vaguely coriaceous; meta-
pleuron duller, closely micropunctate. Pro-
podeal dorsum opaque, with median longi-
tudinal carina; propodeal side dull, closely
obliquely microridged, these extending par-
tially onto hindface with polished center.
Gaster polished, tergum VI with triangular
pygidial plate, surface sparsely setose, apex
rounded. Length 2.75 mm.
Discussion.—Ammoplanus clemente is
known only from the female type. The par-
allel-sided U-shaped clypeal notch with a
long basomedian seta immediately distin-
guishes it from females of A. chemehuevi,
A. loti, A. quabajai, A. unami, and A. van-
yumi. Furthermore, the clypeal notch of A.
clemente lacks the basomedian tooth found
in A. quabajai and A. vanyumi. Other im-
portant features of A. clemente are the oval
area at the inner apex of the eye delimited
by a sharp depression, the deep linear de-
pression of the frons, the smooth, polished
mesopleuron, and the obliquely micro-
ridged propodeal side. It is possible that A.
clemente is the female of either A. sechi or
A. tetli, but based on comparisons with
Pate’s descriptions of the two males, this
appears unlikely.
It is impossible to know if A. clemente is
endemic to San Clemente Island. That will
have to await the results of further collect-
VOLUME 99, NUMBER 1
ing and a much needed revision of North
American Ammoplanus.
Etymology.—The species name is based
on the island from which it comes. It is a
noun in apposition.
Type locality.—California, San Clemente
i viorse Canyon; June, 17;) 1978, A. 'S:
Menke, D. R. Miller and R. W. Rust col-
lectors.
LITERATURE CITED
Bohart, R. M. and A. S. Menke. 1976. Sphecid wasps
of the world; a generic revision. University of Cal-
ifornia Press, Berkeley. 695 ps.
Harris, R. A. 1979. A glossary of surface sculpturing.
State of California Occasional Papers Entomology
(28): 1-31.
Krombein, K. V. 1956. Biological and taxonomic
135
notes on the wasps of Lost River State Park, West
Virginia, with additions to the faunal list (Hyme-
noptera, Aculeata). Proceedings of the Entomo-
logical Society Washington 58: 153-161.
Marshakov, V. G. 1979. Fossorial wasps of the genera
Protostigmus Turner and Ammoplanus Giraud
(Hymenoptera, Sphecidae) from Mongolia and
Middle Asia. Insects of Mongolia 6: 362—374.
Pate, V. S. L. 1943. Nearctic Ammoplanus (Hyme-
noptera: Sphecidae: Pemphredonini). Bulletin of
the Southern California Academy Sciences 41:
141-163.
Rust, R., A. Menke, and D. Miller. 1985. A biogeo-
graphic comparison of the bees, sphecid wasps,
and mealybugs of the California Channel Islands
(Hymenoptera, Homoptera), pp. 29-59. In Menke,
A. S. and D. R., Miller, eds., Entomology of the
California Channel Islands. Santa Barbara Muse-
um of Natural History, Santa Barbara. 178 ps. +
8 maps.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 136-142
THE NEW WORLD ALYDINE GENUS STACHYOCNEMUS STAL
(HEMIPTERA: ALYDIDAE)
CARL W. SCHAEFER AND JOSEPH C. SCHAFFNER
(CWS) Department of Ecology and Evolutionary Biology, University of Connecticut,
U-43, Storrs, CT 06269-3043, U.S.A.; (JCS) Department of Entomology, Texas A&M
University, College Station, TX 77843. U.S.A.
Abstract.—Stachyocnemus is redescribed, and its relationships discussed; except per-
haps for Tollius, Stachyocnemus appears to have no close relatives in North America,
Mexico, or Cuba which encompass the known range of the genus. A specimen from Cuba
represents a new country record, and specimens from Connecticut are a new state record.
Of the two taxa proposed in the genus, S. cinereus Fracker is synonymized with S. apicalis
(Dallas), which is redescribed.
Key Words:
Cuba
Stachyocnemus is a genus of small dis-
tinctive alydines. Although not uncommon,
members of the genus live on the ground
(in dry areas) and are not often collected.
The genus is Nearctic and extends into
southern Mexico; it also occurs in Cuba
(this paper). There have been two species
in the genus, Stachyocnemus apicalis (Dal-
las) and S. cinereus Fracker. Differences be-
tween these two are neither great nor con-
sistent. Here we redescribe the genus, syn-
onymize S. cinereus with S. apicalis, and
redescribe the species.
METHODS AND MATERIALS
In addition to the holotype of Stachyoc-
nemus apicalis and the type series of Stach-
yocnemus cinereus (holotype ¢, Ft. Collins,
CO; allotype °, La Salle, CO; paratype 3,
Gunnison, CO), we have examined more
than 80 specimens from the full range of
the genus. Specimens were examined in
more detail from New Mexico (Socorro
Co.), Texas (Brazos Co.), Connecticut
(Barkhamsted, Litchfield Co. [new state
Alydidae, Heteroptera, Stachyocnemus, genital capsule, Nearctic Region,
record]), Oaxaca, Mexico, and Havana,
Cuba [new country record]. These more
closely studied specimens are in the collec-
tion of CWS or the University of Connect-
icut Insect Collection (the Cuban specimen
is from The Natural History Museum, Lon-
don). The type material of S. cinereus was
borrowed from the National Museum of
Natural History, Smithsonian Institution
(Washington, D.C.), and the type of S. ap-
icalis is from The Natural History Museum
(London).
Genus Stachyocnemus Stal
Stachyocnemus Stal, 1870, p. 215 (orig. de-
scription).
Stachyocnemus Stal, 1873, p. 91 (key).
Stachyocnemus; Lethierry and Severin,
1894-eps 113: (cat):
Stachyocnemis (sic!); Van Duzee, 1906, pp.
386-387 (note).
Stachyocnemis (sic!); Banks, 1910, p. 75
(cat.).
Stachyocnemus; Van Duzee, 1917, p. 115
(cat.).
VOLUME 99, NUMBER 1
Stachyocnemus; Fracker, 1918, p. 275 (de-
scription).
Stachyocnemus; Malloch, 1921, p. 54 (tri-
chobothria).
Stachyocnemus; Torre-Bueno, 1925, p. 28
(ecology and habits).
Stachyocnemus; Blatchley, 1926, pp. 268-—
269 (description).
Stachyocnemus; Torre-Bueno, 1940, p. 159
(note).
Stachyocnemus; Schaefer, 1972a, p.
(loss of scent gland apparatus).
Stachyocnemus;, Schaefer, 1972b, p. 817
(loss of scent-gland apparatus, and ecol-
ogy and biology).
Stachyocnemus; Slater, 1974, pp. 162—163
(relationships).
Stachyocnemus; Schaefer, 1980, pp. 116—
118 (male genital capsule).
19
Head: Dorsum ranging from almost flat
to convex, declivent; about as wide as long;
paraclypei free of and projecting beyond
clypeus, pointed; setae on dorsum of head
dark, scattered, conspicuous; interocellar
distance approximately equal to distance
between ocellus and eye. Antenna short,
about half as long as body, with conspicu-
ous dark scattered setae; first antennal seg-
ment surpassing apex of head but shorter in
total length than head; relative length of
segments in ascending order either 2, 3, 4,
1 or 3, 2, 4, 1. Rostrum reaching mesoster-
num; fourth segment only slightly longer
than third; relative lengths of segments in
ascending order either 3, 4, 1, 2 or 3, 4, 2,
1. Thorax: Pronotum wider than long; an-
terior one-third not arising above level of
collar; posterior corners distinctly acumi-
nate but not drawn out into spines; with
small tooth in middle of posterior margin;
usually a darker triangle evident medially
on pronotum with apex directed anteriorly:
setae dark, scattered, conspicuous; pleural
areas of meso- and metathorax minutely
granulate. Scent gland orifice small, incon-
spicuous, peritreme absent. Scutellum lon-
ger than wide; apex more or less truncate;
apex and edges slightly raised. Corium
137
punctate; apical portion not extending along
costal edge of membrane as in other aly-
dines. Hind coxae widely separated; hind
femur weakly incrassate, armed on poste-
rior surface with two rows of spines; hind
tibia straight, without apical spine, but
armed with two parallel rows of spines, one
row larger and more prominent; tibial setae
present, longer than spines; first hind tarsal
segment almost twice as long as combined
length of second and third segments. Ab-
domen: Robust; sternum IV longer than
other sterna. Male genital capsule and par-
amere: Fig. 1.
Type species: Alydus apicalis Dallas.
Stachyocnemus is easily distinguished
from other North American and Mexican
alydines by this combination of characters:
small size (about three-quarters the size of
the others); lack of an apical hind-tibial
spine, of a costal stridulitrum, and of a me-
tathoracic scent gland peritreme; hind tibia
with two rows of spines; and fourth anten-
nal segment shorter than first. Among the
Alydinae, only Stachyocnemus, Euthetus
Dallas, and Tollius Stal lack the scent gland
peritreme; Euthetus is Old World tropical,
and the New World Tollius lacks the two
rows of spines on the hind tibia.
The generic relationships of Stachyoc-
nemus are unclear. Slater (1974) suggested
the genus was “‘more closely related to Pa-
learctic than to Neotropical genera” (p.
162). Torre Bueno (1940) separated Stach-
yocnemus and Tollius as a group of North
American alydines distinct from Alydus Fa-
bricius, Megalotomus Fieber, and Hyaly-
menus Amyot et Serville. Stachyocnemus
and Tollius do somewhat resemble each
other in habitus, both being smaller than
members of the second group, and both
having obsolescent metathoracic scent
gland apparatuses. However, Tollius has,
and Stachyocnemus lacks, the stridulitrum-
plectrum complex and the genital capsule’s
surcapsular spines, apomorphies possessed
also by Alydus and Megalotomus but not
Hyalymenus (Schaefer et al. 1989). And
Hyalymenus and Stachyocnemus both have
138 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 1.
;
Stachyocnemus apicalis; Texas. Genital capsule (A) and paramere (B) of male. (Note: capsules and
parameres of New Mexico and Connecticut specimens, and of the holotype of S. cinereus, are identical to those
figured here.)
a supporting spur extending from the gen-
ital capsule’s infolded ventral rim to the
cuplike sclerite (Schaefer 1980).
In a cladistic analysis of alydid genera,
Li and Zheng (1993) place Riptortus +
Camptopus as the sister clade of Stachyoc-
nemus. Their text states that the three gen-
era share three apomorphies. However, their
matrix indicates that Riptortus Stal lacks
one of these apomorphies and that Camp-
topus Amyot et Serville lacks another. Thus
Stachyocnemus shares one apomorphy with
both (vesica of aedeagus spiralled), one
with Riptortus (apex of paramere hooked),
and one apomorphy with Camptopus (a
conjunctival character). Yet Stachyocnemus
shares all these apomorphies with Mirpe-
rus, considered by Li and Zheng (1993) the
sister clade of Stachyocnemus + (Riptortus
+ Camptopus). Also, Stachyocnemus
shares two exclusive apomorphies with Tol-
lius (absence of metathoracic scent gland
peritreme, metasternum broad).
Riptortus occurs in the Old World trop-
ics; a few species reach into the southern
temperate regions of Asia. Camptopus ap-
pears to have a rather restricted range in
eastern and southern Europe; and Mirperus
Stal is Afrotropical. The range of none of
these genera supports a sister-group rela-
tionship with Stachyocnemus, which ranges
through the Nearctic south of Canada and
into Mexico and Cuba. Tollius is closer to
Stachyocnemus than are other alydid gen-
era, but the evidence is equivocal and the
relationship weak. We agree with Fracker
(1918) and Schaffner (1964), that Stachyoc-
nemus has no close relatives among the
North American or Mexican alydines or
among any others.
Schaffner (1964) noted that the Stach-
yocnemus male’s genital capsule is
‘“‘unique”’ and that it is “suggestive of”’ that
of the Coreinae (p. 96); later (1979, letter
to CWS) he wrote that “‘the margin of the
genital capsule ... is higher and conse-
quently more like that of the Coreinae.”’ In
addition, Schaefer (1980) showed that the
capsules of both groups have a fused cup-
like sclerite and median projection, and an
entire (not excised) ventral rim. Of these
three features, the last is certainly plesio-
morphic, the second (although advanced)
occurs widely elsewhere, and the first is
both plesiomorphic and common. These
similarities do not indicate a relationship of
Stachyocnemus and the Coreinae.
The known range of the genus is from
VOLUME 99, NUMBER 1
Connecticut (this paper) south into Florida
and west into Alberta (Froeschner 1988),
through Mexico as far south and east as Oa-
xaca (Brailovsky and Flores 1979), and also
into Cuba (this paper).
These insects feed upon seeds of a low-
lying legume (M. H. Sweet, personal com-
munication), and are not predaceous, as
Vestal (1913) suggested.
In 1918 Fracker described a new ‘“‘vari-
ety” of Stachyocnemus apicalis (Dallas),
hitherto the only species in the genus. He
characterized S. apicalis var. cinereus as
paler than the nominate form, more west-
ern, and differing in several structural fea-
tures. Influenced by Fracker’s descriptions
and figures, Torre Bueno (1940) raised var.
cinereus to species rank; later (1941) he
gave the range of each as overlapping in the
Rocky Mountain states.
After examining 78 specimens of both
forms, Schaffner (1964) concluded ‘‘There
is a great range of variability among spec-
imens of the same series”’ and “Characters
given by Fracker (1918) and Torre-Bueno
(1950[sic]) for separation of varietal or spe-
cific designations are of no value’’; and he
synonymized S. cinereus Fracker with S.
apicalis (Dallas). This conclusion, and the
synonymy, were never published.
Fracker (1918) distinguished S. apicalis
from the variety cinereus on the basis of
four characteristics:
@ Juga projecting, or not, from tylus. The
clypeus (tylus) and paraclypei (juga) are
the same in all specimens we have ex-
amined, including the holotype of the S.
apicalis (Florida), a male from Cuba, the
type series of S. cinereus (Colorado), and
specimens from nearby New Mexico. In
the New Mexican specimens, however, a
heavy white pubescence (which is darker
and less dense in all other specimens) ob-
scures the clypeus and paraclypei and
thus their relationship (not so in the S.
cinereus type series or the Cuban speci-
men).
® Body color of S. cinereus gray-flaves-
139
cent, with fine gray pubescence; S. api-
calis black with ferrugineous markings
and little or no gray pubescence. The for-
mer is true (see above) of the New Mex-
ico specimens; the others are reddish
brown (Texas, Oaxaca) or dark brown
(Connecticut). Oddly, the S. cinereus
type series itself is pale brown to tan, and
the holotype of S. apicalis is brown to
dark brown.
@ Head depressed (S. apicalis) or not (S.
cinereus). The head is equally depressed
in all specimens.
@ Medial tooth on pronotum’s posterior
border smaller in S. apicalis than in S.
cinereus, this is true of both holotypes.
This tooth in the Connecticut specimens
and the one from Cuba is smaller than in
the others. But the Oaxaca specimens,
where the tooth is larger, otherwise agree
with Fracker’s description of S. apicalis,
not of S. cinereus; and the tooth of the
New Mexico specimens is smaller than
that of the S. cinereus type series. This
character cuts across Fracker’s descrip-
tions of the two species.
In addition, the pre-antennal head of the
members of the S. cinereus type series, and
of the S. apicalis holotype, is broader—the
paraclypei narrow less sharply—than is the
head of the Connecticut and New Mexico
specimens. But specimens from other parts
of the United States, including Texas, and
from Oaxaca and Cuba, are intermediate.
The New Mexico and Texas specimens
are smaller than the others, as are the male
holotype and paratype of S. cinereus and
the male holotype of S. apicalis. But the
width-length ratios (width across humeri
and total body length) are not greatly dif-
ferent. New Mexico, 0.273; Connecticut,
0.310; Oaxaca, 0.300; Texas, 0.290; S. ci-
nereus holotype, 0.298; allotype, 0.269;
paratype, 0.300; S. apicalis holotype, 0.293;
Cuba 0.301. Similarly, the width-length ra-
tios of the genital capsules from the Texas
(0.800), Connecticut (0.820), and New
Mexico (0.833) specimens differ but little.
140 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
The width-length ratio of the S. cinereus
holotype, however, is 1.0, but in structure it
is identical with that of a Texas specimen
(Fig. 1). The overall size differences there-
fore appear to have resulted from changes
in absolute, not in relative, growth and are
therefore unlikely to be genetic.
The only character which appears to sep-
arate the two forms is the paler body color
and whitish pubescence of S. cinereus. This
is insufficient to warrant distinguishing the
two as species, especially as the genital cap-
sule and parameres of all specimens are
identical (Fig. 1).
Moreover, the distributions do more than
overlap; Stachyocnemus cinereus is an is-
land in a sea of S. apicalis. Fracker (1918)
lists the former from Colorado (type local-
ity), Indiana, Montana, and Arizona; and
we have specimens that fit this description
from New Mexico. S. apicalis is found
throughout North America (Froeschner
1988) and Brailovsky and Flores (1979) list
it through Mexico to Oaxaca; these latter
authors’ description of their specimens fits
S. apicalis better than it does S. cinereus.
For all these reasons, we agree with
Schaffner’s unpublished dissertation (1964),
and synonymize Stachyocnemus cinereus
Fracker with S. apicalis (Dallas).
Stachyocnemus apicalis (Dallas)
Alydus apicalis Dallas, 1852, p. 479 (orig.
description).
Stachyocnemus apicalis;, Stal, 1870, p. 215
(cat.).
Alydus apicalis,; Walker, 1871, p. 159 (cat.).
Alydus apicalis; Walker, 1873, p. 42 (cat.).
Stachyocnemus apicalis;,; Uhler, 1886, p. 12
(list).
Stachyocnemis apicalis; Lethierry and Sev-
erin, 1894, p. 112 (cat.).
Stachyocnemus apicalis; Heidemann,
1902a, p. 32 (distribution).
Stachyocnemus apicalis; Heidemann,
1902b p. 81 (distribution).
Stachyocnemus apicalis; Van Duzee, 1909,
p. 160 (note).
Stachyocnemis (sic!) apicalis; Banks, 1910,
p75 (cak.):
Stachyocnemus apicalis; Fracker, 1918, pp.
275-276, pl. 11, figs. 13, 14 (description,
adult fig.).
Stachyocnemus apicalis; var. apicalis;
Fracker,, 1918, p: 276, pl ohily thea
(note, adult fig.).
Stachyocnemus apicalis; var. cinereus
Fracker, 1918, p. 276, pl. 11, fig. 14 (orig.
description, adult fig.) (New synonymy).
Stachiocnemus (sic!) apicalis; Johnson and
Ledig, 1918, p. 4 (list).
Stachyocnemus apicalis; Malloch, 1921, p.
81, fig. 16 (trichobothria, fig.).
Stachyocnemus apicalis; Blatchley, 1926,
pp. 269-270 (description, note).
Stachyocnemus apicalis var. cinereus;
Blatchley, 1926, p. 270 (note).
Stachyocnemus apicalis; Leonard, 1928, p.
88 (list).
Stachyocnemus apicalis;
1940, p. 159 (note).
Stachyocnemus cinereus; Torre-Bueno,
1940, p. 159 (new status).
Stachyocnemus_ apicalis;
1941 p. 88 (distribution).
Stachyocnemus cinereus; Torre-Bueno,
1942, p. 180 (distribution).
Stachyocnemus apicalis; Sherman, 1948, p.
16 (distribution).
Stachyocnemis (sic!) apicalis; Strickland,
1953, p. 196 (distribution).
Stachyocnemus apicalis; Froeschner, 1988,
p. 8 (cat., distribution).
Stachyocnemus cinereus; Froeschner, 1988,
p. 8 (cat., distribution).
Torre-Bueno,
Torre-Bueno,
Type data: Holotype, male, “‘St. John’s
Bluff, East Florida.”? United States. On de-
posit in The Natural History Museum, Lon-
don.
Description: Pale to dark brown, paler
specimens more heavily beset with pale re-
cumbent setae. Entire insect except for
fourth antennal segment and tarsal seg-
ments usually with conspicuous, decum-
bent, flat almost scale-like microtrichia,
these sometimes silvery under high illumi-
VOLUME 99, NUMBER 1
nation. Side of head with pale line running
from gula onto lateral edge of pronotum.
Pronotum with large triangular region me-
dially, this lightly setose and therefore ap-
pearing darker than lateral regions, these
more heavily beset with pale setae; collar
yellow. Corium sometimes speckled or
lightly mottled with darker brown; if with
pale recumbent hairs, these more dense
along veins. Membrane pale to dark brown,
with dark brown speckling; veins at very
base dark brown, rest of membrane pale
brown. Legs heavily mottled with brown;
spines of femora and tibiae dark brown.
Abdominal venter pale to dark brown, often
paler medially, heavily speckled with
brown or reddish spots; with erect brown
setae interspersed among white recumbent
setae, venter not heavily hirsute; each seg-
ment laterally with large pale spot sur-
rounding spiracle, and with another sublat-
eral spot anterior to trichobothria (latter
spots sometimes obscure or poorly defined).
Total length: 7-9 mm.
NEw DISTRIBUTIONAL RECORDS
CUBA: Grillo Ravelo (1989) listed the al-
ydine genera Burtinus, Hyalymenus, and
Megalotomus from Cuba. A male in The
Natural History Museum (London) of
Stachyocnemus apicalis adds a fourth ge-
nus to the list. Labels: Havana III. 4.v.12/
AWJ Pomeroy collector/U.S. America
A.W.J. Pomeroy. 1919-269/Pres. By Imp.
Bur. Ent. [last label pinned face down].
U.S.A., Connecticut: Although Stachyoc-
nemus apicalis has been recorded from as
close as New York and New Jersey
(Froeschner 1988), it has not been listed
from Connecticut; we list here two males
from Barkhamsted, in the northwestern
part of the state, collected in different
summer months. Labels: CT: Litchfield
Co., Barkhamsted, Farmington River
across from Rt 181, June 11, 1991/R. J.
Packauskas collector [in collection of
C.W.S.]. Second specimen: Barkhamsted
meekRt 318. Litchfield Co., CY Aug. 1,
1991, R. J. Packauskas [fin Univ
Insect Collection].
ACKNOWLEDGMENTS
We thank T. J. Henry (Systematic Ento-
mology Laboratory, U.S.D.A.) for the loan
of the type series of Stachyocnemus ciner-
eus, J. Margerison-Knight (The Natural
History Museum, London) for the loan of
the holotype of Stachyocnemus apicalis and
other specimens, Jane O’Donnell (Univer-
sity of Connecticut Insect Collection,
Storrs) for the loan of specimens, and M. J.
Spring for the figures. We are deeply grate-
ful to Zheng Le-yi and his colleagues at
Nankai University, Tianjin, for providing an
English translation of Li and Zheng (1993),
and to R. C. Froeschner and T. J. Henry,
for comments and additions which have
markedly improved the paper.
LITERATURE CITED
Banks, N. 1910. Catalogue of the Nearctic Hemiptera-
Heteroptera. Philadelphia, American Entomologi-
cal Society.
Blatchley, W. S. 1926. Heteroptera, or true bugs, of
eastern North America. Indianapolis, The Nature
Publishing Company.
Brailovsky, H. and R. Z. Flores. 1979. Contribucion
al estudio de los Hemiptera-Heteroptera de Mé-
xico: XVII. Revision de la familia Alydidae
Amyot y Serville. Anales del Instituto de Biologia
Universidad Nacional Aut6noma México, Series
Zoologia 30: 255-339.
Dallas, W. S. 1852. List of the Specimens of Hemip-
terous Insects in the Collection of the British Mu-
seum. Part II. London, British Museum.
Fracker, S. B. 1918. The Alydinae of the United
States. Annals of the Entomological Society of
America 11: 255-282.
Froeschner, R. C. 1988. Family Alydidae Amyot and
Serville, 1843. The broad-headed bugs, pp. 4—11.
In Henry, T. J. and R. C. Froeschner, eds., Catalog
of the Heteroptera, or True Bugs, of Canada and
the Continental United States. Leiden, E. J. Brill.
Grillo Ravelo, H. 1989. Burtinus notatipennis Stal
(Heteroptera: Alydidae) en Cuba. Communica-
ciones Breves (Revista Centro Agricola) 16(1):
90-92.
Heidemann, O. 1902a. [exhibition note] Proceedings
of the Entomological Society of Washington 5: 32.
. 1902b. Hemiptera, pp. 80—83. Jn Caudell, A.
N., Some insects from the summit of Pike’s Peak,
142 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
found on snow. Proceedings of the Entomological
Society of Washington 5: 74-83.
Johnson, C. and R. Ledig. 1918. Tentative list of He-
miptera from the Claremont-Laguna region. Jour-
nal of Entomology and Zoology 10: 308.
Leonard, M. D. 1928. A list of the insects of New
York with a list of the spiders and certain other
allied groups. New York (Cornell) Agricultural
Experiment Station Memoir 101.
Lethierry, L. and G. Severin. 1894. Catalogue general
des Hémipteéres. Vol. 2. Bruxelles, KF Hayes.
Li Xin-zheng and Zheng Le-yi. 1993. Preliminary
study on the phylogeny of Alydidae (Hemiptera:
Coreoidea). Acta Zootaxon. Sinica 18: 330-343.
[In Chinese, with English abstract.]
Malloch, J. R. 1921. Systematic notes on Hemiptera
Heteroptera. Bulletin of the Brooklyn Entomolog-
ical Society 16: 54-56.
Schaefer, C. W. 1972a. Clades and grades in the Al-
ydidae. Journal of the Kansas Entomological So-
ciety 45: 135-141.
1972b. Degree of metathoracic scent-gland
development in the trichophorous Heteroptera
(Hemiptera). Annals of the Entomological Society
of America 65: 810-821.
1980. The genital capsule of the Alydidae
(Hemiptera: Heteroptera: Coreoidea). Internation-
al Journal of Insect Morphology and Embryology
9: 107-128.
Schaefer, C. W., J. C. Schaffner, and I. Ahmad. 1989.
The Alydus-group, with notes on the alydine gen-
ital capsule (Hemiptera: Heteroptera: Alydidae).
Annals of the Entomological Society of America
82: 500-507.
Schaffner, J.C. 1964. A taxonomic revision of certain
genera of the tribe Alydini (Heteroptera: Corei-
dae). PhD dissertation, Iowa State University,
Ames.
Sherman, F 1948. Coreidae of South Carolina in com-
parison with North Carolina (Hemiptera). Ento-
mological News 59: 15-17.
Slater, J. A. 1974. A preliminary analysis of the der-
ivation of the Heteroptera fauna of the northeast-
ern United States with special reference to the fau-
na of Connecticut. 25th Anniversary Memoirs of
the Connecticut Entomological Society, pp. 145—
PiNN3).
Stal, C. 1870. Enumeratio hemipterorum. Part 1. Kon-
gliga Svenska Vetenskaps-Akademiens Handlin-
gar 9: 1-232.
1873. Enumeratio Hemipterorum. Part 3.
Kongliga Svenska Vetenskaps-Akademiens Hand-
lingar 11: 1—163.
Strickland, E. M. 1953. An annotated list of the He-
miptera (s.l.) of Alberta. The Canadian Entomol-
ogist 85: 193-214.
Torre-Bueno, J. R. de la. 1925. Methods of collecting,
mounting and preserving Hemiptera. Canadian
Entomologist 57: 6—10, 27—32, 54-57.
Torre-Bueno, J. R. de la. 1940. Tollius vanduzeei n.
sp., with notes on the genera Tollius Stal and
Stachyocnemus Stal (Heteroptera, Alydidae). Bul-
letin of the Brooklyn Entomological Society 35:
159-161.
1941. A synopsis of the Hemiptera-Heter-
optera of American north of Mexico. Entomolo-
gica Americana 21: 41-122.
1942. Notes on Coreidae. Bulletin of the
Brooklyn Entomological Society 37: 180.
Uhler, P. R. 1886. Checklist of the Hemiptera Heter-
optera of North America. Brooklyn, Brooklyn En-
tomological Society.
Van Duzee, E. P. 1906. New North American Heter-
optera. Entomological News 17: 384-391.
1909. Observations on some Hemiptera
taken in Florida in the spring of 1908. Bulletin of
the Buffalo Society of Natural Sciences 9: 149—
230.
. 1917. Catalogue of the Hemiptera of Amer-
ica north of Mexico, excepting the Aphididae,
Coccidae and Aleurodidae. University of Califor-
nia Publications on Entomology 2, 902 pp.
Vestal, A. G. 1913. An associational study of Illinois
sand prairie. Bulletin of the Illinois State Labo-
ratory of Natural History 10: 1-96.
Walker, EF 1871. Catalogue of the Specimens of He-
miptera Heteroptera in the Collection of the Brit-
ish Museum. London, E. Newman.
. 1873. Catalogue of Hemiptera Heteroptera in
the British Museum. Supplement London, E. W.
Janson.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 143-155
PARASITOIDS (HYMENOPTERA: BRACONIDAE: APHIDIINAE) OF APHIDS
ON BIG SAGEBRUSH (ARTEMISIA TRIDENTATA NUTTALL) AND PRAIRIE
SAGE (ARTEMISIA LUDOVICIANA NUTTALL) IN WASHINGTON STATE
K. S. PIKE, P STARY, D. ALLISON, G. GRAF, L. BOYDSTON, AND T. MILLER
(KSP) Entomologist and (DA, GG, LB) Research Technologists, Washington State Uni-
versity, Irrigated Agriculture Research and Extension Center, 24106 N Bunn Road, Pros-
ser, WA, 99350 U.S.A.; (PS) Entomologist, Institute of Entomology, Academy of Sciences
of the Czech Republic, BraniSovska 31, 370 05 Ceské Budéjovice, Czech Republic; (TM)
Insectary Manager, Northwest Biocontrol Insectary and Quarantine, Washington State
University, Pullman, WA 99164-6382, U.S.A.
Abstract.—Parasitoid-aphid associations on big sagebrush, Artemisia tridentata Nuttall
and prairie sage, Artemisia ludoviciana Nuttall in Washington State are determined, and
the parasitoid species are reviewed and keyed. Two new species of aphidiine parasitoids
are described and illustrated: Praon artemisicola Pike and Stary, n. sp. (hosts: Epa-
meibaphis atricornis Gillette & Palmer, Flabellomicrosiphum sp. and Pseudoepameiba-
phis tridentatae Wilson; also from mixed populations of Flabellomicrosiphum knowltoni
Smith, Obtusicauda filifoliae Gillette and Palmer, and Pleotrichophorus sp. on A. triden-
tata), and Trioxys artemisiarum Pike and Stary, n. sp. (host: Macrosiphoniella ludo-
vicianae (Oestlund) on A. ludoviciana). The sagebrush areas of the region are relatively
widespread with mainly indigenous flora; the associated aphids and parasitoids are also
indigenous and constitute a relatively rich diversity of species. Twelve species of sage-
brush and prairie sage aphid-parasitoids are known from the region.
Key Words: sagebrush, Artemisia, aphids, parasitoids, Praon artemisicola n. sp., Trioxys
artemisiarum n. sp., biodiversity
Big sagebrush (Artemisia tridentata Nut-
tall) (Fig. 1) occurs across vast reaches and
fragmented tracts of land within the interior
areas of the Pacific Northwest where pre-
cipitation is commonly less than 50 cm per
year. Big sagebrush is indigenous to the re-
gion and somewhat competitive with peren-
nial grasses (Whitson et al. 1991) and other
plants. The sagebrush communities com-
monly adjoin agricultural lands, sometimes
dryland grain fields, sometimes irrigated
fields. Prairie sage (Artemisia ludoviciana
Nuttall) is also common in the region, but
restricted more to streambanks and flood
plains.
A recent review of Pacific Northwest
aphidiine parasitoids by the authors (Pike et
al. 1996) indicated a number of plant com-
munities in which more detailed research
should be undertaken, including sagebrush.
Further, a catalog by Marsh (1979) of North
American aphidiines showed little infor-
mation for the Northwest. We theorized that
sagebrush or sagebrush and other plants in
the sagebrush ecosystem, especially in un-
disturbed settings, might harbor parasitoids
(Fig. 2) useful against aphids in adjoining
crops or serve as a stable source for such.
Distinguishing tritrophic associations (par-
asitoid-aphid-plant) in ecosystems or agro-
144 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-2. 1, Big sagebrush landscape, WA, Benton Co., Rattlesnake Hills, 1-VI-96 (photo by L. C. Wright,
Res. Tech. Supervisor, Washington State University, Prosser, WA). 2, Obtusicauda coweni aphids on big sage-
brush parasitized by Lysiphlebus utahensis, WA, Yakima Co., near Wenas Lake, 30-V-96 (photo by K. S. Pike).
VOLUME 99, NUMBER |
ecosystems constitute one of the first steps
needed in biodiversity research, and for ad-
vancing the use of or enhancing aphid bi-
ological controls. The purpose of this re-
search was to define the diversity of aphi-
diine parasitoids and their associated aphid-
hosts on big sagebrush and prairie sage,
describe new species as appropriate, and
present a key to the parasitoids of big sage-
brush and prairie sage aphids.
MATERIALS AND METHODS
Big sagebrush and prairie sage were ex-
tensively sampled for aphids and aphid-par-
asitoids across eastern Washington, cover-
ing Asotin, Benton, Douglas, Franklin,
Klickitat, Kittitas, and Yakima counties. A
total of 130 samples was taken over three
years (1994-96), and 831 parasitoids were
reared and examined. The collected live
aphids from which parasitoids were reared,
were held in semi-transparent plastic con-
tainers (300 ml, 10 cm dia X 4 cm ht or
3500 ml, 18 cm dia X 13 cm ht) on clipped
foliage at laboratory temperatures of 20 +
3°C for 25—30 days to provide time for par-
asitoids to develop; adults generally
emerged within 10 days.
New descriptions were based on whole
dry and dissected slide-mounted specimens
evaluated under microscope at 40 to 600.
Measurements were recorded in millime-
ters. Holotypes were dry-mounted on paper
tabs and pinned. Descriptive terminology is
after Huber and Sharkey (1993). Holotypes
are deposited in the National Museum of
Natural History, Smithsonian Institution,
Washington DC (USNM).
REVIEW OF PARASITOID SPECIES OF BIG
SAGEBRUSH (BS) AND PRAIRIE
SAGE (PS) APHIDS
Parasitoid species are listed alphabetical-
ly. References, if any, are listed at the end
of each record of aphid host, location, and
date of collection. All aphids were collected
and identified by the authors, unless other-
wise indicated. Numbers in parentheses
represent authors’ codes of voucher speci-
145
mens (aphid-parasitoids) in Washington
State University (WSU)-Prosser collec-
tions.
Aphidius polygonaphis Fitch, ali from
aphids on PS: USA, WA, Yakima Co.,
Yakama Indian Res., Yakima Chief Rd.,
16-V-96, from Macrosiphoniella ludovici-
anae and Pleotrichophorus sp. (96G053),
Mabton Rd., 25-IV-96 (96K004), and
Lower Mill Cr. Rd., 25-V-96, (96K054)
from Macrosiphoniella ludovicianae.
Binodoxys coruscanigrans (Gahan): USA,
WA, Yakima Co., Mabton Grade, 23-V-
96, and near Wenas Lake, 30-V-96, from
Obtusicauda coweni on BS (96G068,
96G115).
Binodoxys clydesmithi Pike & Stary, all
from aphids on BS: USA, WA, Franklin
Co., Kahlotus Hwy, Neff Farm, 28-V-96,
from Obtusicauda artemisicola (96G096);
Kittitas Co., Lost Lake, 7-VII-94, from
Obtusicauda coweni and Zyxaphis sp.
(det. G. Remaudiére) (94K042) (Pike et
al. 1996).
Ephedrus californicus Baker, all from
aphids on BS unless indicated otherwise:
USA, WA, Klickitat Co., Roosevelt, 9 &
24-IV-96, from Obtusicauda _ sp.
(96G011, 96GO15), 16-IV-96, from Ob-
tusicauda artemisiphila (96GO12); Yaki-
ma Co., Mabton Grade, 14-V-96, from
Obtusicauda coweni (96G026); Yakama
Indian Res., Mabton Rd., 25-IV-96, from
Macrosiphoniella nr. ludovicianae on PS
(96K004), 16-V-96, from Macrosiphon-
iella ludovicianae and Pleotrichophorus
pseudoglandulosus on PS (96G049).
Lysaphidus adelocarinus (Smith), all from
aphids on BS: USA, WA, Benton Co.,
Arid Lands Ecol. Reserve, Rattlesnake
Springs, 10-V-96, from Epameibaphis
atricernis, Microsiphoniella sp., Obtusi-
cauda sp. and Pleotrichophorus spp.
(95G032), Snively Springs, 19-V-95,
from Epameibaphis atricornis and Pseu-
doepameibaphis tridentatae (95G072),
near Rothrock and Pearl Rds., 29-VI-95,
from Flabellomicrosiphum knowltoni,
146 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Pseudoepameibaphis tridentatae and Zy-
xaphis canae (95G280); Franklin Co., Ju-
niper Dunes Wilderness, 23-V-95, from
Flabellomicrosiphum knowltoni and Mi-
crosiphoniella acophorum (95G088,
95G089), Kahlotus Hwy, Neff Farm,
28-V-96, from Obtusicauda artemisicola
(96G096); Kittitas Co., Ellensburg,
20-VI-95, from Pseudoepameibaphis tri-
dentatae (95G198), Taneum Rd., 11-IX-
95, from Pleotrichophorus sp., Pseudo-
epameibaphis glauca, and Zyxaphis sp.
(95G586); Klickitat Co., Alderdale Cr. &
Columbia River, 28-IV-95, from Pleotri-
chophorus sp. (95GO12), 2-IV-96, from
Flabellomicrosiphum tridentatae, Obtu-
sicauda sp. and Pseudoepameibaphis tri-
dentatae (96G007), near Alderdale, 9-V-
95, from Pleotrichophorus sp. (95G026),
Flabellomicrosiphum sp., Obtusicauda
filifoliae and Pseudoepameibaphis triden-
tatae (95G027), 11-VII-95, from Flabel-
lomicrosiphum tridentatae and Pseudo-
epameibaphis essigi (95G345), Roose-
velt, 24-IV-96, from Obtusicauda sp.
(96G015); Yakima Co., Moxee Hwy 24,
15-V-95, from Epameibaphis atricornis
and Flabellomicrosiphum knowltoni
(95G066), Plank Rd., 15-V-95, from Pseu-
doepameibaphis tridentatae (95G069),
Fort Simcoe, 30-V-95, from Flabellomi-
crosiphum knowltoni and Pseudoepamei-
baphis tridentatae (95G091), Yakama In-
dian Res., near White Swan, 6-VI-95,
from Obtusicauda filifoliae and Pseudoe-
pameibaphis tridentatae (95G110), Yak-
ama Indian Res. near Satus, 16-V-96,
from Epameibaphis atricornis, Flabel-
lomicrosiphum sp., and Pseudoepamei-
baphis tridentatae (96G062), Yakama In-
dian Res., Mabton Rd., 25-IV-96, from
Obtusicauda sp. (96KO005), Yakima Fir-
ing Range, 24-VII-95, from Obtusicauda
sp., and Pseudoepameibaphis tridentatae
(95G370).
Lysaphidus ramithyrus (Smith): USA, WA,
Kittitas Co., Taneum Rd, 11-[X-95, from
Pleotrichophorus sp. and Zyxaphis sp. on
BS (95G585).
Lysiphlebus utahensis (Smith), all from
aphids on BS: USA, WA, Franklin Co.,
Kahlotus Hwy, Neff Farm, 28-V-96, from
Obtusicauda artemisicola (96G096);
Klickitat Co., near Roosevelt, 2 & 9-IV-
96, from Obtusicauda filifoliae (96GO009)
and Obtusicauda sp. (96G011); Yakima
Co., Mabton Grade, 23-V-96, and near
Wenas Lake, 30-V-96, from Obtusicauda
coweni (96G068, 96G115, 96G122).
Praon artemisaphis Smith, all from aphids
on BS: USA, WA, Klickitat Co., Roose-
velt, 9-IV-96, from Obtusicauda sp.
(96G011); Yakima Co., Mabton Grade,
14 & 23-V-96, and near Wenas Lake,
30-V-96, from Obtusicauda coweni
(96G026, 96G068, 96G115), Yakama In-
dian Res., Mabton Rd., 25-IV-96, from
Flabellomicrosiphum knowltoni and
Pleotrichophorus sp. (96K003), from
Obtusicauda sp. (96KO05).
Praon artemisicola Pike & Stary,
new species
(Figs. 3-8)
Diagnosis.—Following the key by John-
son (1987), the new species has several
characters (radial sector length, distal ab-
scissa of R1 length, and stigma width) in
common with P. artemisaphis Smith, but
differs in the number of antennal segments
(13-14) and status of the m-cu vein (almost
to completely effaced).
Etymology.—The new species name is
derived from its association with big sage-
brush, Artemisia tridentata.
Description.—Female: Head. Malar
space equal to ¥ of eye length. Face gla-
brous, with a row of setae along orbits. An-
tenna 13-14 segmented, short, filiform, as
long as head, mesosoma and basal third of
metasoma together. Flagellomere 1 (= F,)
4X as long as wide, with sparse long
oblique setae slightly longer than half seg-
ment diameter, no longitudinal placodes. F,
3x as long as wide, with one placode. Mid-
dle flagellomeres 2.5X as long as wide,
with sparse oblique setae subequal to seg-
VOLUME 99, NUMBER 1
6
Figs. 3-8.
Praon artemisicola (illustrations not to equal scale). 3, Mesonotum. 4, Genitalia. 5, Ovipositor
sheath. 6, Forewing. 7, Forewing lower margin close-up. 8, Metasomal tergum |. Abbreviations: R1, radial
abscissa (= metacarpus); Rs + M, wing vein.
ment diameter. Preapical flagellomere as
wide as middle flagellomeres.
Mesosoma. Mesonotum (Fig. 3) with all
lobes glabrous, with a few scattered setae
along the notauli. Propodeum smooth.
Forewing (Figs. 6—7). Stigma 4 as long
as wide; distal abscissa of RI (= metacar-
pus) relatively short, equal to ¥, of stigma
length. Radial sector equal to stigma width;
median vein discolored in the proximal por-
tion; m-cu vein effaced except sometimes
for a short discolored proximal stub. Lower
marginal setae (Fig. 7) 5X as long as sur-
face setae.
Legs. Hind femur with sparse semi-erect
setae that are slightly longer than half seg-
ment diameter.
Metasoma. Metasomal tergum | (Fig. 8)
square, with groups of setae close to hind
corners.
Genitalia (Figs. 4—5).
Coloration. Head dark brown, mandibles
brown, palpi light brown. Antenna brown,
pedicel and basal % of F, yellow to light
brown with lighter basal ring. Mesosoma
dark brown. Wings subhyaline, venation
light brown. Fore leg yellow brown to
brown, middle and hind legs brown, base
of tibiae lighter. Metasoma brown, tergum
1 light brown. Ovipositor sheath brown.
Body length about 1.6—1.8 mm.
Male: Antenna 16—17 segmented, long,
subequal to body length. Concolorous,
somewhat darker than female. Antenna
dark brown, narrow apex of pedicel and a
narrow basal ring of F, light brown. Meta-
soma brown, tergum | dark brown.
Holotype [2 ].—USA, WA, Yakima Co.,
Alderdale and Glade Rd., 9-V-95, reared
from (mixed aphids) Epameibaphis atricor-
148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
nis Gillette & Palmer, Flabellomicrosiphum
sp., and Pseudoepameibaphis tridentatae
(Wilson) taken from big sagebrush, Arte-
misia tridentata Nuttall, coll. G. Graf, sam-
ple no. 95G030. Deposited in USNM.
Paratypes.—reared from aphids on big
sagebrush, Artemisia tridentata: USA, WA,
Benton Co., Crosby Rd. 1-V-95, from Pseu-
doepameibaphis tridentatae (95GO015, 3 ¢),
Hwy 240 and Horn Rapids, 12-V-95, from
Pseudoepameibaphis tridentatae (95G041,
1 2); Klickitat Co., Alderdale Rd., 9-V-95,
from Flabellomicrosiphum sp., Pseudoe-
pameibaphis tridentatae, and Obtusicauda
filifoliae (95G0O27, 1 ¢), near Alderdale,
9-V-95, from Pleotrichophorus sp. and
Pseudoepameibaphis tridentatae (95G028,
1 2), Alderdale Rd., 5mi N of Hwy 14,
12-V-95, from Epameibaphis atricornis
(95G037, 1 2); Yakima Co., Alderdale and
Glade Rd., 9-V-95, from Epameibaphis
atricornis, Flabellomicrosiphum sp. and
Pseudoepameibaphis tridentatae (95G030,
2 2), Moxee Hwy, 15-V-95, from Epamei-
baphis atricornis, Flabellomicrosiphum
knowltoni (95G066, 1 2, 2 3), Plank Rd.,
15-V-95, from Pseudoepameibaphis triden-
tatae (95G069, 2 3). Material deposited in
part in the Washington State University
Collection, James Museum, Pullman, WA;
the Washington State University-Prosser
Collection, and the collection of P. Stary,
Ceske Budéjovice, Czech Republic.
Praon occidentale Baker: USA, WA, Ya-
kima Co., Yakama Indian Res., Mabton
Rd., 25-IV-96, from Macrosiphoniella lu-
dovicianae on PS (96KO04).
Trioxys artemisiarum Pike & Stary,
new species
(Figs. 9-12)
Diagnosis.—Following the key by Smith
(1944), the new species has characters in
common with 7. bonnevillensis Smith and
T. infrequens Smith. T. bonnevillensis dif-
fers from the new species in having some-
what shorter prongs, four perpendicular se-
tae dorsally, and with oblique setae distinct-
ly longer than prong diameter. 7. infrequens
(based on Smith’s 1944 original descrip-
tion) differs from the new species in having
prongs fused along basal halves, and fla-
gellomeres approximately equal in length
and unusually long. Also, the antenna of the
new species is 12-segmented vs. 11-seg-
ment in 7. bonnevillensis and T. infrequens.
Etymology.—The new species name is
derived from its association with Artemisia.
Description. Female: Head. Eyes promi-
nent, strongly convergent to clypeus. Malar
space equal to \ of eye length. Tentoriocu-
lar line equal to % of intertentorial line. An-
tenna 12-segmented, filiform, reaching to
middle of metasoma; middle to apical fla-
gellomeres equal in width, gradually shorter
towards apex of flagellum. Second flagel-
lomere (F,) 3X as long as wide, middle fla-
gellomeres 2.3X as long as wide.
Mesosoma. Mesonotum with simple
rows of relatively sparse setae along effaced
notauli of disc. Propodeum (Fig. 11)
smooth, with two short divergent carinae in
distal portion.
Forewing (Fig. 9) stigma triangular, al-
most 3X greatest width; distal abscissa of
R1 (= metacarpus) relatively short, shorter
than % stigma length or distinctly shorter
than stigma width. Radial sector short,
somewhat longer than stigma width. Hind
femur and tibia with adpressed setae.
Metasoma. Metasomal tergum | (Fig. 10)
somewhat longer than 2 spiracular width,
dilating from poorly prominent spiracular
tubercles to apex, about 1.5X as wide at
apex as across spiracles.
Genitalia (Fig. 12). Ovipositor sheath
distinctly claw-shaped. Prongs long and
straight except for upward curve at apex,
distinctly separated along their length, 7—8
long perpendicular setae on dorsal surface
(setae gradually increasing in length to-
wards apex of prong, length 2—3X adjacent
prong diameter), numerous shorter oblique
setae on ventral surface (length subequal to
adjacent prong diameter), sternal base of
prongs with a single row of long, longitu-
dinal setae (subequal in length to dorsal se-
VOLUME 99, NUMBER 1
Figs. 9-12.
tergum 1. 11, Propodeum. 12, Genitalia. Abbreviation: st, spiracular tubercle.
tae), and prong apex with two longitudi-
nally dilated setae.
Coloration. Distinctly bicolorous, with
some variation. Head yellow; ocellar trian-
gle brown, sometimes brown from frons to
vertex. Antenna dark brown, most of scape
and pedicel, and whole of F, yellow. Me-
sosoma brown black, sometimes prothorax
yellow brown. Tegulae yellow. Wings sub-
hyaline, venation brown. Legs brown, tinge
darkest in hind leg; trochanter, base and
narrow apex of tibia and tarsomere | or
most of tarsus yellow. Metasoma distinctly
bicolorous, middle part prevalently dark
brown; tergum | and central spot at tergum
2 light brown. Middle terga sometimes with
149
eS ~
Trioxys artemisiarum (illustrations not to equal scale). 9, Forewing, in part. 10, Metasomal
light lateral spots; apex yellow. Last tergum
with central brown spot, apical half of ovi-
positor sheath brown; sternum and prongs
yellow.
Body length 2.3—2.5 mm.
Male: Antenna 14-segmented. Coloration
generally dark brown; apex of pedicel and
mandibles yellow. Metasomal tergum 1 and
base of tergum 2 light brown.
Holotype [2].—USA, WA, Asotin Co.,
Heller’s Bar, 20-VI-95, reared from Macro-
siphoniella ludovicianae (Oestlund) taken
from big sagebrush, Artemisia tridentata
Nutall, coll. T. Miller, sample no. 95T146.
Deposited in USNM.
Paratypes.—USA, WA, Asotin Co.,
150 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Couse Creek and Heller’s Bar, 20-VI-95,
from Macrosiphoniella ludovicianae on BS
(951126, 2°93 2.6: 951 146,°2""6)): "and
1'82VII-95"'on' PS (951257; 2 “2, 39d) We=
posited in part in the Washington State Uni-
versity Collection, James Museum, Pull-
man, WA; the Washington State University-
Prosser Collection, and the collection of P.
Stary, Ceske Bud&éjovice, Czech Republic.
Trioxys bonnevillensis Smith, all from
aphids on BS unless indicated otherwise:
USA, WA, Benton Co., Arid Lands Ecol.
Reserve, Rattlesnake Springs, 10 -V-95,
from Epameibaphis atricornis, Microsi-
phoniella sp., Obtusicauda sp., and Pleo-
trichophorus spp. (95G032), 19-V-95,
from Epameibaphis atricornis and Pseu-
doepameibaphis tridentatae (95GO072),
8-VI-95, Epameibaphis sp. and Pseudo-
epameibaphis tridentatae (95G151),
Bennett Springs, 2-V-95, from Epamei-
baphis atricornis, Pleotrichophorus sp.,
and Pseudoepameibaphis tridentatae
(95G018), Hwy 14, 12-V-95, from Pseu-
doepameibaphis tridentatae (95G039),
Prosser, 8-VI-95, from Pseudoepameiba-
phis tridentatae (95K032), Rotha Rd,
2-V-95, from Obtusicauda artemisiae
(95G017), Rothrock Rd., 29-VI-95, from
Flabellomicrosiphum knowltoni, Pseu-
doepameibaphis tridentatae, and Zyxa-
phis canae (95G280); Douglas Co.,
Badger Mtn., 13-VIH-95, from Zyxaphis
canae Williams, coll. R. L. Gillespie
(95B012); Franklin Co., Juniper Dunes
Wilderness, 23-V-95, from Flabellomi-
crosiphum tridentatae and Microsiphon-
iella acophorum (95G088, 95G089); Kit-
titas Co., Ellensburg, 20-VI-95, from Pseu-
doepameibaphis tridentatae (95G198),
Klickitat Co., Alderdale, 11-VII-95, from
Flabellomicrosiphum tridentatae and
Pseudoepameibaphis essigi (95G345),
Alderdale Cr., 28-IV-95, from Flabello-
microsiphum tridentatae (95GO013), from
Epameibaphis utahensis, Obtusicauda fil-
ifoliae and Pseudoepameibaphis triden-
tatae (95G014), 9-V-95, from Flabello-
microsiphum sp., Obtusicauda filifoliae
and Pseudoepameibaphis tridentatae
(95G027), 12-V-95, from Pseudoepamei-
baphis tridentatae (95GO038), Roosevelt,
2-IV-96, from Obtusicauda filifoliae
(96GO09), 9-IV-96, Obtusicauda sp.
(96G011), 16-IV-96, from Obtusicauda
artemisiphila (96G0O12); Yakima Co.,
Hwy 82, 12-V-95, from Epameibaphis
atricornis and Pseudoepameibaphis tri-
dentatae (95G043), 12-V-95, from Pseu-
doepameibaphis tridentatae (95G044),
Moxee Hwy, 15-V-95, from Epameiba-
phis atricornis and Flabellomicrosiphum
knowltoni (95G066), Hwy 82, 15-V-95,
from Epameibaphis atricornis and Pseu-
doepameibaphis tridentatae (95G067),
Fort Simcoe, 30-V-95, from Flabellomi-
crosiphum knowltoni and Pseudoepamei-
baphis tridentatae (95G091), White
Swan, 6-6-95, from Obtusicauda filifol-
iae and Pseudoepameibaphis tridentatae
(95G110), Yakima Firing Range, 24-VII-
95, from Obtusicauda sp. and Pseudo-
epameibaphis tridentatae (95G370), Yak-
ama Indian Res., Chief Rd, 25-IV-96,
from Obtusicauda sp. (96KO005), from
Obtusicauda sp. and Pseudoepameiba-
phis tridentatae on PS (96G020), Yaka-
ma Indian Res. near Satus, 16-V-96, from
Epameibaphis atricornis, Pseudoepamei-
baphis tridentatae and Flabellomicrosi-
phum sp. (96G062).
KEY TO PARASITOID SPECIES (FEMALES)
REARED FROM APHIDS ON ARTEMISIA SPP.
IN EASTERN WASHINGTON STATE
1. Forewing Rs + M vein present, sometimes
more or less colorless, but distinct in fore
part (Bigss6; 14> 225523) Seis. eee 2
— Forewing Rs + M vein absent (Figs. 13,
725) hae etre Petpet wbrney A, cededete ts IE rove 0c 5
2(1). Forewing 1R1 and 1 + 2Rs cells separate;
rein \esinl jornenrernte (Weis, M4) wee bags e
aie hip APiogewewehe Sayre Ephedrus californicus Baker
— Forewing I1R1 and 1 + Rs confluent; r-m
vein absent (Praon) (Figs. 6, 22, 23) .... 3
3(2). Forewing m-cu vein present (Figs. 22, 23);
antenna’ I6—1'8*seemented’ == 725-0. =: +
— Forewing m-cu vein absent (Fig. 6); anten-
VOLUME 99, NUMBER 1
4(3).
S\(1l)}
8(7).
na 13-14 segmented
. Praon artemisicola Pike & Stary, n. sp.
Forewing distal abscissa of Rl (= meta-
carpus) and radial sector short, equal to
width of stigma (Fig. 22); antenna 16—17
segmented Praon artemisaphis Smith
Forewing distal abscissa of R1 and radial
sector long, at least 1.5—2.0 times longer
than width of stigma (Fig. 23); antenna 17—
18 segmented Praon occidentale Baker
Forewing r-m vein present (Figs. 13, 24,
25); last abdominal sternite simple, without
prongs; ovipositor sheath slightly curved
po wanls (Were, IO, 17, 22) Geco scenes o~ 6
Forewing r-m vein absent (Figs. 27, 28);
last abdominal sternite with fork-like ac-
cessory prong; ovipositor sheath curved
downward (Figs. 12, 30, 32, 34) ....... 9
. Forewing M + m-cu vein complete (Fig.
13) Aphidius polygonaphis Fitch
Forewing M + m-cu vein incomplete (Fig.
DS) se Bice Roe ROC Lae ee 7
. Propodeum with more or less distinct are-
ola or at least carinae (Figs. 19—20); meta-
somal tergum 1 somewhat parallel-sided
(nicer) Uaysaphidits) yess 8
Propodeum smooth (Fig. 18); metasomal
tergum 1 somewhat triangular, distinctly
enlarged at apex (Fig. 15)
Lysiphlebus utahensis (Smith)
Propodeum with two divergent carinae in
distal portion (length of carinae variable)
(Fig. 20) .. Lysaphidus adelocarinus (Smith)
Propodeum areolated, with small central
pentagonal areola (size and shape variable)
(Fig. 19) ....Lysaphidus ramithyrus (Smith)
. Metasomal tergum 1 with spiracular and
secondary tubercles (Fig. 26)
Metasomal tergum | with spiracular tuber-
cles only (Figs. 10, 31)
al
. Lower apical part of metasoma yellow,
prongs dark; spiracular and secondary tu-
bercles of metasomal tergum 1 of equal
size, distinctly separated from each other;
prongs feeble, with 4—5 long setae on dor-
sal surface in basal portion (Fig. 33); ovi-
positor sheath broad in basal two-thirds, al-
most claw-shaped (Fig. 32)
Sa eens Binodoxys coruscanigrans (Gahan)
Lower apical part of metasoma brown,
concolorus with remaining metasoma,
prongs dark brown; spiracular and second-
ary tubercles of metasomal tergum | close-
ly spaced, secondary tubercles slightly
greater than spiracular tubercles (Fig. 26);
Ovipositor sheath not claw-shaped, com-
paratively narrow at base (Fig. 30); prongs
strong, with 7 long setae on dorsal surface
11(9).
(BS,
151
(Fig. 30); stigma length 3x width (Fig. 28)
aio ROR Binodoxys clydesmithi Pike & Stary
Prongs of last metasomal sternum short, ar-
cuate, each bearing 4 long perpendicular
setae on dorsal surface (Fig. 34); radial
sector shorter than stigma width (Fig. 27)
Ov EC-5. 0 ERENCES Trioxys bonnevillensis Smith
Prongs of last metasomal sternum long,
straight, with apices curved upward, each
bearing 7—8 perpendicular setae on dorsal
surface (Fig. 12); radial sector longer than
stigma width (Fig. 9)
... Trioxys artemisiarum Pike & Stary, n. sp.
APHID-PARASITOID-PLANT
ASSOCIATIONS LISTING
big sagebrush; PS, prairie sage. *, par-
asitoid reared from a collection of
mixed aphids; specific association
shown considered valid by authors)
Epameibaphis atricornis Gillette &
Palmer
*T ysaphidus adelocarinus (BS)
*Praon artemisicola (BS)
*Trioxys bonnevillensis (BS)
Epameibaphis utahensis Knowlton &
Smith
*Trioxys bonnevillensis (BS)
Flabellomicrosiphum knowltoni Smith
Fl
*Lysaphidus adelocarinus (BS)
Praon artemisaphis (BS)
*Praon artemisicola (BS)
*Trioxys bonnevillensis (BS)
abellomicrosiphum tridentatae (Wil-
son)
*T ysaphidus adelocarinus (BS)
*Trioxys bonnevillensis (BS)
Flabellomicrosiphum sp.
*T ysaphidus adelocarinus (BS)
*Praon artemisicola (BS)
*Trioxys bonnevillensis (BS)
Macrosiphoniella ludovicianae (Oes-
tlund)
* A phidius polygonaphis (PS)
Ephedrus californicus (PS)
*Praon occidentale (PS)
*Trioxys artemisiarum (BS)
Microsiphoniella acophorum (Smith &
Knowlton)
*T ysaphidus adelocarinus (BS)
152 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 13-17. Various features of parasitoids of big sagebrush and prairie sage aphids (illustrations not to
equal scale). 13, Aphidius polygonaphis, forewing. 14, Ephedrus californicus, forewing. 15-17, Lysiphlebus
utahensis. 15, Metasomal tergum 1. 16, Genitalia. 17, Ovipositor sheath. Abbreviations: M + m-cu, Rs + M
and r-m are wing veins; 1M, IRI, | + 2Rs are wing cells.
*Trioxys bonnevillensis (BS) *Binodoxys clydesmithi (BS)
Microsiphoniella sp. *Lysiphlebus utahensis (BS)
*Lysaphidus adelocarinus (BS) *Lysaphidus adelocarinus (BS)
*Trioxys bonnevillensis (BS) Obtusicauda artemisiphila (Knowlton
Obtusicauda artemisiae (Cowen ex Gil- and Allen)
lette & Palmer) *Ephedrus californicus (PS)
*Trioxys bonnevillensis (BS) Trioxys bonnevillensis (BS)
Obtusicauda artemisicola (Williams) Obtusicauda coweni Hunter
VOLUME 99, NUMBER 1 153
Figs. 18-25. Various features of parasitoids of big sagebrush and prairie sage aphids (illustrations not to
equal scale). 18, 24, Lysiphlebus utahensis. 18, Propodeum. 24, Forewing, in part. 19, 21, Lysaphidus ramithyrus.
19, Propodeum. 21, Metasomal tergum 1. 20, 25, Lysaphidus adelocarinus. 20, Propodeum. 25, Forewing. 22,
Praon artemisaphis, forewing. 23, Praon occidentale, forewing. Abbreviations: R1 (radial abscissa, = metacar-
pus), Rs (radial sector), Rs+M, M+m-—cu, m-cu, and r—m are wing veins; 1M, wing cell.
* Binodoxys clydesmithi (BS) Lysaphidus adelocarinus (BS)
* Binodoxys coruscanigrans (BS) *Lysiphlebus utahensis (BS)
*Ephedrus californicus (BS) Praon artemisicola (BS)
*Lysiphlebus utahensis (BS) Trioxys bonnevillensis (BS)
*Praon artemisaphis (BS) Obtusicauda sp.
Obtusicauda filifoliae (Gillette & Palmer) *Ephedrus californicus (BS)
*Ephedrus californicus (PS) Lysaphidus adelocarinus (BS)
154 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 26-34. Various features of parasitoids of big sagebrush and prairie sage aphids (illustrations not to
equal scale). 26, 28, 30, Binodoxys clydesmithi. 26, Metasomal tergum |. 28, Forewing, in part. 30, Genitalia.
27, 31, 34, Trioxys bonnevillensis. 27, Forewing, in part. 31, Metasomal tergum 1. 34, Genitalia. 29, Lysaphidus
adelocarinus, genitalia. 32, 33, Binodoxys coruscanigrans. 32, Ovipositor sheath. 33, Prong. Abbreviations: se,
secondary tubercle; st, spiracular tubercle.
*Praon artemisaphis (BS) Pseudoepameibaphis essigi Knowlton &
Trioxys bonnevillensis (BS) Smith
Pleotrichophorus pseudoglandulosus *Tysaphidus adelocarinus (BS)
(Palmer) *Trioxys bonnevillensis (BS)
Ephedrus californicus (PS) Pseudoepameibaphis glauca Gillette &
Pleotrichophorus sp. Palmer
*Lysaphidus adelocarinus (BS) *Lysaphidus adelocarinus (BS)
*Lysaphidus ramithyrus (BS) Pseudoepameibaphis tridentatae (Wil-
Praon artemisaphis (BS) son)
*Praon artemisicola (BS) *Lysaphidus adelocarinus (BS)
*Trioxys bonnevillensis (BS) *Praon artemisicola (BS)
VOLUME 99, NUMBER 1
*Trioxys bonnevillensis (BS)
Zyxaphis canae Williams
*Lysaphidus adelocarinus (BS)
*Trioxys bonnevillensis (BS)
Zyxaphis sp.
*Binodoxys clydesmithi (BS)
*Lysaphidus adelocarinus (BS)
*Lysaphidus ramithyrus (BS)
The aphid-parasitoid fauna on big sage-
brush and prairie sage is indigenous and rel-
atively diverse: in total 12 species of para-
sitoids were found associated with at least
15 species of aphids. A part of these para-
sitoid species are distributed over other
western states and Mexico (Marsh 1979,
Stary and Remaudiere 1982, 1983, Stary
1983, Johnson 1987). Three of the parasit-
oid species (Aphidius polygonaphis, Ephed-
rus californicus, and Praon occidentale) are
known to attack selected pestiferous aphids
in adjoining crops or forest (Pike et al.
1996). The extent of their value, however,
in terms of possible positive connections or
movement between sage plant communities
(endemic, typically undisturbed ecosys-
tems) and neighboring agroecosystems or
introduced landscapes is still largely unde-
termined.
ACKNOWLEDGMENT
We express thanks to the Yakama Indian
Nation for permission to collect and eval-
uate aphid and aphid parasitoids on their
lands, and G. Remaudiere (Muséum Na-
tional d’ Histoire Naturelle, Paris) for valu-
able help in aphid identification. This work
was made possible in part by funds provid-
ed by Washington State University and the
U.S. Department of Agriculture.
LITERATURE CITED
Huber, J. T. and M. J. Sharkey. 1993. Structure, pp.
13—59: In Goulet H: and J. T. Huber, eds., Hy-
menoptera of the World: An Identification Guide
to Families. Research Branch, Agriculture Cana-
da, Ottawa, Ontario, Publication 1894/E, 668 pp.
Johnson, J. W. 1987. A revision of the species of
Praon Haliday in North America north of Mexico
(Hymenoptera: Aphidiidae). Canadian Entomolo-
gist 119: 999-1025.
Marsh, P. M. 1979. Aphidiidae, pp. 295-313. In
Krombein, K. V., P. D. Hurd Jr., D. R. Smith, and
D. B. Burks, eds., Catalog of Hymenoptera of
America North of Mexico. Vol. 1, Symphyta and
Apocrita (Parasitica). Smithsonian Institution
Press, Washington D.C., 1198 pp.
Pike, K. S., P. Stary, R. Miller, D. Allison, L. Boyd-
ston, G. Graf, and T. Miller. 1996. New species
and host records of aphid parasitoids (Hymenop-
tera, Braconidae, Aphidiinae) from the Pacific
Northwest, USA. Proceedings of the Entomolog-
ical Society of Washington 98: 570-591.
Smith, C. EF 1944. The Aphidiinae of North America
(Braconidae: Hymenoptera). Ohio State Univer-
sity Contributions in Zoology and Entomology
No. 6, 154 pp.
Stary, P. 1983. New species and records of aphid par-
asitoids from Mexico (Hymenoptera, Aphidiidae).
Acta Entomologica Bohemoslovaca 80: 35—48.
Stary, P. and G. Remaudiére. 1982. New genera, spe-
cies and host records of aphid parasites (Hyme-
noptera: Aphidiidae) from Mexico. Annales de la
Societé Entomologique de France, N.S., 18: 107—
Ie
1983. Complements to the aphid parasitoid
fauna of Mexico (Hymenoptera, Aphidiidae). An-
nales de la Societé Entomologique de France,
N.S., 19: 113-116.
Whitson, T. D., L. C. Burrill, S. A. Dewey, D. W. Cud-
ney, B. E. Nelson, R. D. Lee, and R. Parker. 1991.
Weeds of the west. Western Society of Weed Sci-
ence, Western U.S. Land Grant Universities Co-
operative Extension Services, and University of
Wyoming, Pioneer of Jackson Hole, Jackson, WY,
630 pp.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 156-161
A NEW SPECIES OF THE NEOTROPICAL GENUS LISSOSCARTA STAL
(HOMOPTERA: CICADELLIDAE: CICADELLINAE) THAT MIMICS WASPS
GABRIEL MEJDALANI AND MARCIO FELIX
Departamento de Zoologia, Instituto de Biologia, Universidade Federal do Rio de Ja-
neiro, Caixa Postal 68044, 21944-970 Rio de Janeiro RJ, Brasil.
Abstract.—Lissoscarta beckeri, new species, a leafhopper that mimics wasps, is de-
scribed from the states of Rond6nia and Mato Grosso, Brazil. The new species can be
distinguished from the other species of the genus by the following features: males with
an elongate dorsoapical process in pygofer; aedeagus with a conspicuous dorsal lobe and
without processes; paraphyses arched dorsally; abdominal sternum VII of females with a
concavity on posterior margin. All members of the genus Lissoscarta have a strong con-
striction at the base of the abdomen, the forewings completely hyaline, and the form of
the pronotum similar to the mesoscutum of a wasp. The size and color are similar to those
of wasps of the tribe Polybiini (Vespidae: Polistinae). Morphological comparisons and
field observations suggested that at least six species in four different genera of that tribe
could be models for L. beckeri.
Key Words:
Six valid species of the Neotropical ge-
nus Lissoscarta Stal were recorded by
Young (1977). The type species of the ge-
nus, L. vespiformis (Fabricius, 1803), has
morphological and behavioral characters
that enable it to mimic wasps (Evans 1947,
Boulard 1978). The resemblance between
L. vespiformis and wasps was first noted by
Fabricius (1803) in the original description
of the species (as Cicada vespiformis). The
abdomen is strongly constricted at the base,
the forewings are completely hyaline, and
the form of the pronotum is similar to the
mesoscutum of a wasp. When threatened,
this leafhopper spreads its wings in a sim-
ilar way to that of a wasp in the resting
position, showing the constriction at the
base of the abdomen. This behavior, in
which the mimic suddenly exposes charac-
teristics that resemble those of its model, is
called ‘‘ostensible mimicry’? (Boulard
1978). It is not found in any other known
Cicadellidae, Lissoscarta, new species, mimicry, wasps
leafhopper genus. All the remaining species
of Lissoscarta have the morphological fea-
tures just mentioned (Young 1977), sug-
gesting that “‘ostensible mimicry”’ is a char-
acteristic of the genus. Unfortunately, no
behavioral data are available for these other
species.
In the present paper a new species of Lis-
soscarta from Brazil is described. Morpho-
logical comparisons, as well as field obser-
vations (J. Becker, personal communica-
tion), suggested that wasps of the tribe Po-
lybiini (Vespidae, Polistinae) could be
models for the new species. The characters
supporting this view are discussed below.
Acronyms for collections in which the
specimens used in this study are deposited
are as follows: DZUP (Departamento de
Zoologia da Universidade Federal do Pa-
rana, Curitiba, Brazil), MNRJ (Museu Na-
cional do Rio de Janeiro, Rio de Janeiro,
Brazil), and MZSP (Museu de Zoologia da
VOLUME 99, NUMBER 1
157
Figs. 1-12.
4, Right hindwing. 5—9, Male genitalia. 5, Pygofer and subgenital plate, lateral view. 6, Subgenital plate, ventral
view. 7, Right style and connective, dorsal view. 8, Aedeagus, lateral view. 9, Paraphyses, dorsal view. 10-12,
Female. 10, Abdominal sternum VII, ventral view. 11, Pygofer, lateral view. 12, Second valvula of ovipositor,
lateral view.
Universidade de Sao Paulo, Sao Paulo, Bra-
zil). In quotations of label data, a virgule
(/) separates lines on a label and a semi-
colon separates information on different la-
bels. Morphological terminology follows
mainly Young (1977), excepting that of
wing veins, which follows Dworakowska
(1988). Nomenclature of wasp species fol-
Lissoscarta beckeri. 1, Head and thorax, dorsal view. 2, Head, lateral view. 3, Right forewing.
lows Richards (1978) with modifications
introduced by Carpenter and Day (1988).
Lissoscarta beckeri Mejdalani and Felix,
new species
(Figs. 1-13)
Diagnosis.—Lissoscarta beckeri can be
distinguished from the other species of the
158 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
genus by the following features: males with
dorsoapical margin of pygofer with elon-
gate process extending ventrally; aedeagus
with conspicuous dorsal lobe, without pro-
cesses; rami of paraphyses arched dorsally;
females with abdominal sternum VII with
median concavity on posterior margin.
Description.—Length of male 14.8—16.0
mm; female 16.6—16.8 mm. Surface of pro-
notum convex in dorsal view, with sulcus
near anterior margin. Forewing (Fig. 3)
elongate, completely hyaline. Hindwing
(Fig. 4) strongly narrowed, with slightly en-
larged jugal lobe, vein AP’ greatly reduced
and AP” + JA absent. Abdomen (Fig. 13)
strongly constricted at base between seg-
ments III and IV. Remaining morphological
characters of head and thorax as in the ge-
neric description of Young (1977: 148).
Color: Crown, pronotum and scutellum
yellow. Crown (Fig. 1) with inconspicuous
transverse brown line at apex of coronal su-
ture, anterior margin with pair of black
spots, posterior margin with sinuate trans-
verse black stripe. Pronotum (Fig. 1) with
sinuate transverse black stripe on anterior
margin, median and posterior portions with
elongate brown spot on each side of larger
triangular brown spot, lateral margins with
elongate brown spot. Mesonotum (Fig. 1)
with pair of conspicuous black maculae ex-
tending from base to scutellar suture, basi-
lateral margins with small black spot, scu-
tellum with transverse dark brown stripe.
Forewing (Fig. 3) with fumose area extend-
ing from near base of anal margin caudo-
laterally across clavus and becoming dif-
fuse in corium. Hindwing with most of ScP
+ R + MA yellow. Face yellow. Clypeus
(Fig. 2) with lateral and ventral margins
black and with pair of longitudinal black
stripes on muscle impressions extending to
apical spots of crown. Clypellus (Fig. 2)
with blackish apex. Lora (Fig. 2) with black
spot on inferior portion. Genae (Fig. 2) with
black spot below eyes, dorsoposterior mar-
gin black. Thoracic sterna yellow. Anterior
tibiae brown, with black stripe along ventral
margin. Abdomen (Fig. 13) with tergites
yellowish-brown, posterior margins with
transverse yellow stripe.
Male genitalia (Figs. 5—9): Pygofer (Fig.
5) moderately produced in lateral view,
with posterior margin convex; with macro-
setae numerous, lacking only on basidorsal
area; dorsoapical margin with elongate pro-
cess extending ventrally. Subgenital plates
(Figs. 5, 6) in ventral view elongate, trian-
gular, extending posteriorly almost as far as
pygofer apex; with uniseriate macrosetae;
microsetae also present. Style (Fig. 7) ex-
tending posteriorly considerably beyond
apex of connective, without preapical lobe;
preapical area with group of setae; apex
rounded. Connective (Fig. 7) T-shaped. Ae-
deagus (Fig. 8) symmetrical, with conspic-
uous dorsal lobe in lateral view; ventral
margin with longitudinal sulcus. Paraphyses
(Fig. 9) with pair of asymmetrical rami di-
rected dorsally.
Female genitalia (Figs. 10—12): Abdom-
inal sternum VII (Fig. 10) broad in ventral
view, with median concavity on posterior
margin. Pygofer (Fig. 11) well produced in
lateral view, triangular, with apex narrowly
rounded; most macrosetae on apex and ex-
tending anteriorly along ventral margin,
small number below dorsal margin. Second
valvulae of ovipositor (Fig. 12) expanded
beyond basal curvature, preapical promi-
nence discrete, apex acute; shaft bearing
teeth throughout expanded portion, except
on apical area, teeth quadrate, sloping and
bearing minute secondaries, apex with nu-
merous denticles on dorsal and ventral mar-
gins.
Type material.—Holotype: 6, Brazil,
“Ouro Preto/ d’Oeste-RO/ 21-X-1986/ J.
Becker col.”, MNRJ. Paratypes: One 4,
one 2, same data as holotype, MNRJ. One
3, ‘“‘Pimenta Bueno-RO/ 23-X-1986/ J.
Becker col.”’, MNRJ. One ¢d, one @, “‘Vila
Vera MT/ Brasil X-1973/ M. Alvarenga
Ker ss DZUP:
Additional material.—Dr. M. W. Nielson
(Brigham Young University, Utah, U.S.A.)
identified, using a manuscript of our paper,
eight specimens of L. beckeri in the collec-
VOLUME 99, NUMBER 1
tion of the Utah State University. He has
kindly sent us the information on the labels
of these specimens: ““One 6, one °, Brazil:
Rondonia, 62 mi. SE Ariquemes, 15-22
March 1991, W. Hanson, G. Bohart; two d,
one ¢, same data as above except 13-25
April 1992, W. J. Hanson; one 6, two 2,
same data as above except 17-24 May
1989, 180 m, W. J. Hanson.”
Notes.—This species keys to Lissoscarta
vespiformis in Young’s (1977) key. The
convex posterior margin of the pygofer
(Fig. 5) and the conspicuous dorsal aedea-
gal lobe (Fig. 8) of L. beckeri are similar to
those of L. vespiformis, but the latter does
not have pygofer processes and the rami of
its paraphyses are more symmetrical; the
posterior margin of female abdominal ster-
num VII (Fig. 10) is distinctly concave in
L. beckeri and only slightly emarginate in
L. vespiformis. The unpaired dorsal aedea-
gal processes of L. schlingeri Young, L.
pereneensis Young, L. catutara Young, and
L. nipata Young will readily distinguish
them from L. beckeri; in L. catutara the
posterior margin of female sternum VII is
also only slightly emarginate. Lissoscarta
pebasensis Young is known only from the
holotype female (Young 1977); the abdom-
inal sternum VII of this species has a reg-
ularly convex posterior margin that distin-
guishes it from L. beckeri. The new species
is named in honor of Prof. Johann Becker
(Museu Nacional, Rio de Janeiro), who has
collected many interesting Homoptera in
Brazil.
DISCUSSION
We have studied wasp species belonging
to four genera of the tribe Polybiini: Age-
laia Lepeletier [A. fulvofasciata (Degeer),
A. flavipennis (Ducke), and A. hamiltoni
(Richards)], Pseudopolybia Saussure [P.
difficilis (Ducke)], Polybia Lepeletier [P.
(Pedothoeca) emaciata Lucas], and Mis-
chocyttarus Saussure [M. (Haplometrobius)
undulatus (Ducke)]. The specimens exam-
ined are deposited either in MNRJ or MZSP
and were determined by O. W. Richards,
159
excepting a single specimen of A. fulvofas-
ciata from the type locality of L. beckeri,
which was identified by the authors. These
species are very similar in terms of color
pattern, a fact indicating that they are part
of a group of Miillerian mimics. The head
and thorax in all of them are yellow, being
similarly marked by dark spots and stripes.
The abdominal tergites are yellowish-
brown and often present a yellow transverse
stripe at posterior margin.
The features mentioned above, which are
here exemplified by A. fulvofasciata (Fig.
14), are mimicked by L. beckeri (Fig. 13).
This represents a case of Batesian mimicry.
The presence of dark spots and stripes on
the head and thorax of the leafhopper, as
well as the transverse yellow stripes at the
posterior margin of the abdominal tergites,
are remarkable characters. All wasps stud-
ied occur within the geographical range of
L. beckeri (see Richards 1978). They are
approximately the same size as the mimic.
In terms of morphology, the constriction
at the base of the abdomen of L. beckeri is
an important character (Fig. 13). This con-
striction mimics the constricted zone (peti-
ole) at the base of the abdomen of the
wasps. The fore- (Fig. 3) and hindwings
(Fig. 4) of L. beckeri are hyaline, resem-
bling those of the polybiines mentioned.
The hindwings are also strongly narrowed,
having a form very different from that usu-
ally found in other members of the subfam-
ily Cicadellinae. The vein AP’ is greatly re-
duced and AP” + JA is absent. The convex
pronotum of L. beckeri mimics the meso-
scutum of the wasps.
Agelaia fulvofasciata (Degeer) is a model
for L. beckeri in the type locality of the
latter. It is apparently more common than
the leafhopper (J. Becker, personal com-
munication). Becker also observed that L.
beckeri can rest upon the leaves with its
wings spread in wasp-like manner. This be-
havior is similar to the “bluffing display”
described by Boulard (1978) for L. vespi-
formis. In agreement with its wide range,
commonness and aggressive behavior, A.
160 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 13-14.
soscarta beckeri, dorsal habitus of female.
fulvofasciata is mimicked by many species
of wasps and other insects (Richards 1978).
Indeed, Richards and Richards (1951) ob-
served that it is also a model for L. vespi-
formis.
Using Vane-Wright’s (1976) terminology
and analytic schemes, this case of mimicry
would be Class VI (antergic defensive).
This class includes Bates’ original formu-
lation of mimicry (Vane-Wright 1976). The
predators (operators) from which such
mimicry affords protection are unknown.
Therefore, it is not possible to establish
whether this mimicry is disjunct (mimic,
model, and operator are different species)
or bipolar (model and operator are the
same). More field observations are neces-
sary to settle this question. At least two oth-
er cases of antergic defensive mimicry have
13, Lissoscarta beckeri, dorsal habitus of female. 14, Agelaia fulvofasciata, a model of Lis-
been reported in Homoptera. Hogue (1984)
speculated that lanternflies (Fulgora spp.)
avoid predation by mimicking arboreal liz-
ards. Zolnerowich (1992) described a
nymph of Amycle sp. (Fulgoridae) that
mimics jumping spiders (Salticidae). In
these two cases the model and operator
were considered the same (bipolar).
ACKNOWLEDGMENTS
We are grateful to J. Becker (MNRJ) for
making available for study specimens under
his care and for providing invaluable infor-
mation about mimicry in L. beckeri. We
also thank the following persons for lending
specimens of Lissoscarta: R. R. Cavichioli
(DZUP), M. D. Webb (The Natural History
Museum, London), and U. R. Martins
(MZSP). C. R. Brandéo (MZSP) helped a
VOLUME 99, NUMBER I
great deal with the identification of A. ful-
vofasciata and made available for study
wasps under his care. The habitus drawings
of L. beckeri and A. fulvofasciata were
inked by M. EF Pess6éa. The manuscript ben-
efited from the comments of J. Becker, EF
A. Bockmann, A. L. Carvalho, C. H. Die-
trich, L. EF Dorvillé, W. J. Knight, J. L. Nes-
simian, M. W. Nielson, E. R. da-Silva, M.
D. Webb, and two anonymous reviewers.
LITERATURE CITED
Boulard, M. 1978. Premier cas de “‘mimétisme osten-
sible’? chez les Homopteres Auchénorhynques
(Insecta). Compte Rendu Hebdomadaire des Sé-
ances de |’ Academie des Sciences 287 (D): 1389—
1S9ie
Carpenter, J. M. and M. C. Day. 1988. Nomenclatural
notes on Polistinae (Hymenoptera: Vespidae). Pro-
ceedings of the Entomological Society of Wash-
ington 90: 323-328.
Dworakowska, I. 1988. Main veins of the wings of
Auchenorrhyncha (Insecta, Rhynchota: Hemely-
trata). Entomologische Abhandlungen aus dem
Staatlichen Museum fiir Tierkunde in Dresden 52:
63-108.
Evans, J. W. 1947. A natural classification of leaf-
161
hoppers (Jassoidea, Homoptera). Part 3: Jassidae.
Transactions of the Royal Entomological Society
of London 98: 105—271.
Fabricius, J. C. 1803. Systema Rhyngotorum secun-
dum Ordines, Genera, Species adiectis Synony-
mis, Locis, Observationibus, Descriptionibus.
Apud Carolum Reichard. Brunswick. 314 pp.
Hogue, C. L. 1984. Observations on the plant hosts
and possible mimicry models of “Lantern Bugs”
(Fulgora spp.) (Homoptera: Fulgoridae). Revista
de Biologia Tropical 32: 145-150.
Richards, O. W. 1978. The social wasps of the Amer-
icas excluding the Vespinae. British Museum
(Natural History). London. 580 pp. + 4 pls.
Richards, O. W. and M. J. Richards. 1951. Observa-
tions on the social wasps of South America (Hy-
menoptera Vespidae). Transactions of the Royal
Entomological Society of London 102: 1-169 +
I-IV pls.
Vane-Wright, R. I. 1976. A unified classification of
mimetic resemblances. Biological Journal of the
Linnean Society of London 8: 25-56.
Young, D. A. 1977. Taxonomic study of the Cicadel-
linae (Homoptera: Cicadellidae). Part 2. New
World Cicadellini and the genus Cicadella. North
Carolina Agricultural Experiment Station Techni-
cal Bulletin 239: 1-1135.
Zolnerowich, G. 1992. A unique Amycle nymph (Ho-
moptera: Fulgoridae) that mimics jumping spiders
(Araneae: Salticidae). Journal of the New York
Entomological Society 100: 498-502.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 162-170
BIONOMICS OF LASIOGLOSSUM (EVYLAEUS) MATIANENSE (BLUTHGEN)
(HYMENOPTERA: HALICTIDAE), THE PREDOMINANT POLLINATING BEE
IN ORCHARDS AT HIGH ALTITUDE IN THE GREAT HIMALAYA OF
GARHWAL, U.P., INDIA
SUZANNE W. T. BATRA
Bee Research Laboratory, Bldg. 476, U.S. Department of Agriculture, Beltsville, MD
20705; U:S:A;
Abstract—The halictine bee, Lasioglossum (Evylaeus) matianense (Bliithgen) is the
most abundant bee in early May, pollinating apples soon after snowmelt at Harsil, India
(elevation 2600 m), and pollinating Prunus sp. at Gangotri, India (elevation 3100 m). It
nests in south-facing, sunny banks, where density reaches 2403 nests per m? (X:290 nests
per m2). The nests of this probably solitary species are short, sinuous, nearly horizontal
burrows, ending in 1 or 2 cells at a mean depth of 6.2 cm. This bee thermoregulates its
nests by placing them in relatively cool, moist soil among grass roots near the tops of
the banks.
Key Words:
Worldwide, there are about 2,000 species
of halictine bees, popularly called ‘sweat
bees’ because some lick salty sweat. They
are important pollinators of crops and wild
plants in all terrestrial habitats, including
Arctic tundra, the world’s highest moun-
tains, wettest rainforests, driest deserts, vast
prairies, temperate forests, and urban gar-
dens. Because most species are small, dark,
and inconspicuous, and nest underground,
people often overlook them, even when
they are very abundant in habitats. They are
also difficult to identify taxonomically. Hal-
ictine bees, including Evylaeus, are of great
interest for the study of sociobiology and
evolution, because they have such a diver-
sity of social behavior, ranging from strictly
solitary species to communal, semisocial,
and eusocial (sensu Batra 1966) bees that
share nests (see Packer and Knerer 1985 for
a review). Some species may be eusocial in
a favorable habitat with a long growing sea-
son, but live a solitary life where the grow-
Himalaya, pollination, apple, Prunus, bee, nests, thermoregulation
ing season is too short to permit the devel-
opment of social colonies, for example, L.
(Evylaeus) calceatum (Scopoli) (Sakagami
and Munakata 1972).
The subgenus Evylaeus of Lasioglossum
is most abundant and diverse in cold cli-
mates of North America and Eurasia. Two
species, L. (E.) rufitarse (Zetterstedt) and L.
(E.) borealis Svensson, Ebmer, Sakagami,
are Holarctic (Sakagami and Toda 1986).
Evylaeus is divided into two groups, those
in which the females have a carina on the
posterior edge of the propodeum (Svensson
et al. 1977), and those without a carina.
Carinate species often construct their nests
so that the subterranean brood cells are ar-
ranged to form delicate earthen combs that
are surrounded by airfilled cavities (Batra
1990). In general, nests of carinaless spe-
cies lack combs (based on a sample of few-
er than 20 species). According to A. W. Eb-
mer (in litt.), L. (Evylaeus) matianense
(Bliithgen, 1926) belongs to the holarctic L.
VOLUME 99, NUMBER 1
(E.) nitidiusculum taxonomic species-
group.
From March 5 to May 8, 1995, I under-
took a survey of bees that pollinate rosa-
ceous fruit crops in the Garhwal Himalaya,
U.P, India (Batra in press). The Shivalik
range and Lesser Himalaya were surveyed
at several locations (77°30' to 79°00'E;
30°15’ to 30°30'N), at altitudes from 579 to
3100 m above M.S.L. during March and
April. Beginning on May 1, I studied the
pollinators of apples growing at and near
Harsil (elevation 2600 m; 78°45’E,
31°02'N) and Prunus sp. at Gangotri (ele-
vation 3100 m; 78°57’E, 31°00'N). This
area is in the subalpine zone of the Great
Himalaya Range (Mani 1962; Mani 1978),
about 20 km south of the Tibet (China) bor-
der, in a deep valley on the upper Bhagir-
athi Ganga River, surrounded by snow-cap-
ped peaks of over 6000 m elevation. Apples
are grown on terraced slopes to 3100 m el-
evation. Timberline is at about 3600 m.
Mornings were clear, but strong southerly
orographic updrafts brought haze, clouds
and chilly precipitation after noon, and
cold, katabatic drafts blew down from the
glaciers to the north at night. The pollinat-
ing bees on early-bloom apples and full-
bloom pears at first included no honey bees.
Almost all of the bees were halictines, es-
pecially L. (E.) matianense. Various Diptera
were abundant on the apple flowers. No
Apis cerana EF live at, or north of, the apple-
growing district of Sukhi (elevation 2487
m) where, according to local growers, bee-
keeping is not practiced. Apis laboriosa
Sm. foragers suddenly appeared on apple
blossoms at Harsil on May 5. They were
some 600 km west of the previously known
range of this seasonally migrating giant
honey bee in Nepal (for behavioral details,
see Batra 1996).
Although some 9000 species of seed
plants grow in the 1450 km? area of the
western Himalaya that is above 1200 m el-
evation (Polunin and Stainton 1984), and
some 4000 species are endemic to the Hi-
malaya (Mani 1978), very little is known
163
about their pollinators, most of which may
be endemic, as are other Himalayan insects
(Mani 1962). Due to its remoteness and dif-
ficulty of access, the bees of the Great Hi-
malaya Range are practically unknown. No
work on the pollination of the temperate
crops that are grown in these enormous, ex-
tensive, and rugged mountains has been
conducted. This is significant, because this
region is adjacent to the centers of origin
for several important crops, thus, efficient
coevolved pollinators of these crops should
occur there. Previous work on crop polli-
nation in the Himalaya has been conducted
in the Lesser Himalaya, which are influ-
enced by the monsoon and have a more
moderate climate (Batra in press).
The Harsil area is of particular ecological
interest because it lies just north of the
crestline of the Great Himalaya Range. Har-
sil and Gangotri are in the rain shadow of
Srikanta (6132 m elevation), Jaonli (6632
m) and Phating Pithwara (6904 m), which
partially block the southeast monsoon. Ar-
temisia and other xerophytes that are char-
acteristic of Middle Asia (Mani 1978) grow
there. The river valley and its orchards are
surrounded by extensive glaciated areas.
Apple orchards were first planted beside the
river at Harsil about 75 years ago, and some
of the original trees are still living there,
according to local growers. Apples and oth-
er fruit trees were subsequently planted ex-
tensively on terraces where the native de-
odar cedars were cleared, from Sukhi to the
south (2500 m elevation) and at several lo-
cations along the river, to the east and up-
stream of Harsil, as far as Gangotri (ele-
vation 3100 m). These may be among the
world’s higher orchards (Fig. 2). Apple va-
rieties include c.v. ‘Delicious’, c.v. ‘Bijou’
and a wild-type, seedy, ‘Kashmiri’ apple.
Other fruits grown at sheltered sites along
the river in the area include pear, plum,
quince, chulu apricot, and almonds.
POLLINATION
I collected apple pollinators on May |
and from May 5 to 7 at Harsil, by sweeping
164 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 1.
Location where L. (E.) matianense was collected in Garhwal: H, Harsil; G, Gangotri. The stippled
area represents the Great Himalaya Range (Himadri), where many peaks are over 6,000 m and valleys are over
2000 m. Other locations are: T, Tibet; N, Nepal; U, Uttarkashi; D, Delhi. Shimla (S) and Mussoorie (M) are
‘hill stations’ on the crests of ridges at 2206 m and 2006 m respectively, in the Lesser Himalaya (Himanchal).
Between Himanchal and Himadri are subtropical valleys, as low as 500 m in elevation.
the canopy with a long-handled insect net.
Although sunrise occurred at 6:30 local
time, due to the chilly nights, bees did not
begin to fly until 9:00, and maximum for-
aging activity was between 10:00 and 13:
OO. In the early afternoon, strong southerly
winds and cloudiness developed daily. The
bees were very sensitive to any dimming of
the brilliant sunshine, and they left the or-
chard before rain, hail, and snow fell.
The most abundant pollinating bee on
apple bloom at Harsil was L. (E.) mati-
anense. On May 1, 93% (98 of 106) of the
bees that were collected were this species.
At another site near Harsil on May 6, 75%
(49 of 65 bees) on apples were L. (E.) ma-
tianense. I also collected pollinators on
Prunus sp. (? P. mira) growing near melt-
ing snowbanks on May 3 and 4 at Gangotri
(apples were not yet in bloom there). At
this location, also, L. matianense was the
predominant bee. For comparison, at
VOLUME 99, NUMBER 1
Fig. 2.
Map of the Harsil area, showing surrounding high peaks (numbers indicate elevation in meters),
glaciated areas (stippled), and locations of the orchards (hatched areas). S = Sukhi; H = Harsil; G = Gangotri;
T = Tibet.
Dhanaulti (2200 m elevation) in the Lesser
Himalaya, Apis cerana were by far the
most abundant pollinators of apples in
mid-April; beekeeping being widely prac-
ticed there. Halictine bees, especially both
sexes of L. (E.) marginatum Brullé, were
second in abundance (at about 10% of the
total). The predominance of halictines at
high elevations, and the scarcity of An-
drena, Osmia, and Bombus, both in num-
bers of individuals and in diversity of spe-
cies, were unexpected findings. These
three genera (especially Andrena spp.) are
the most important native, unmanaged ap-
ple pollinators in North America and Eu-
rope (Boyle and Philogéne 1983; Boyle-
166 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Makowski 1987; Scott-Dupree and Win-
ston 1987, and references therein).
AGGREGATION AND THERMOREGULATION
A large aggregation of the nests of L. (E.)
matianense was discovered on May 7, in a
steep, sunny, south-facing slope, about 100
m above the orchard near Wilson Cottage
at Harsil. Foraging females were returning
with large loads of apple pollen. This slope
was an overgrazed pasture, where most of
the original deodar cedars had been re-
moved. Scattered large trees provided
patches of shade. Several trails had been cut
into the slope by grazing yaks, mules, and
goats, and the bees’ nests were in the ca.
0.5 to 1.0 m-high bank of soil that had been
exposed on the upslope sides of these live-
stock trails. The underlying soil was glacial
till, a mixture of fine, micaceous silt and
sand, mixed with pebbles, cobbles, and flat,
gleaming, micaceous rocks. Above it was
20-30 cm layer of moist loam, filled with
the rootlets of short grasses and forbs, such
as Taraxacum and Fragaria, that were just
beginning their spring growth. This area
was probably glaciated within the past
10,000 years. The Bhagirathi Ganga River
emerges from a retreating glacier near Gan-
gotri at about 4000 m elevation. Next to the
zigzag livestock trails, L. (E.) matianense
nests formed patches of dense aggregations
where the south-southeast aspect and inso-
lation appeared to be favorable. Counts of
all nest entrances in 31 meter-square sec-
tions were made at 5 locations within a 100
m? area, along 4 of the trails. Total nests
per m? included samples with mostly soil,
and samples that contained large, obstruct-
ing rocks and roots, which were less favor-
able for nesting. Results are as follows:
Site 1 (7 m? samples): 20 (rocky) to 348
nests per m*; X = 165 nests per m7’.
Site 2 (6 m?* samples): 10 (roots) to 545
nests per m?; xX = 115 nests per m?.
Site 3 (12 m?* samples): 50 to 2403 nests
per m?; X = 376 nests per m?.
Site 4 (5 m?* samples): 15 (rocky) to 369
nests per m*; X = 182 nests per m’.
Site 5 (1 m? sample): 130 nests.
The mean for all 5 sites was 290 nests
per m*; 7303 nest entrances were counted
within the 31 m7? total area surveyed. Be-
cause similarly suitable habitat extended for
an estimated 300 m along the livestock
trails on the slope above the orchard, as
many as 87,000 nests of L. (E.) matianense
may have been present.
The Himalaya are notable for extreme
variability of microclimates, the result of
their high elevation, rugged terrain, conti-
nentality, and relatively low latitude. A
combination of diurnal insolation and noc-
turnal radiation, atmospheric aridity, cold,
low oxygen pressure, rapid desiccation,
strong winds, and extreme daily tempera-
ture fluctuations affects all life. Small, mi-
croclimatic differences in aspect, light and
shade may change the length of the growing
season and time of erop maturity at a lo-
cation by up to 3 weeks (Whiteman 1985).
At the elevation of Harsil, the frost-free
growing season is about 24 weeks (Mani
1978). In the Great Himalaya, the atmo-
spheric mean temperature increases rapidly
from March to June, but then it stabilizes
by July, due to monsoon cloud cover, be-
fore declining after August (Mani 1978).
Lasioglossum (E.) matianense has exploited
the brief period of intense insolation and
warmth that exists in May and June, when
it makes and provisions its nests and its
brood develops. It has also exploited the
phenomenon that south-facing slopes re-
ceive twice as much solar radiation as
north-facing slopes (Whiteman 1985), by
nesting in south-facing slopes. In North
America, the vernal bee L. (Evylaeus) com-
agenensis (Knerer and Atwood) similarly
nests where it maximizes insolation (Batra
1990a), as do Andrena alleghaniensis Vi-
ereck (Batra 1990b) and A. fenningeri Vi-
ereck (Batra, in litt.). In Japan, L. (Evy-
laeus) duplex (Dalla Torre) selects nest sites
VOLUME 99, NUMBER 1
that receive direct morning sunshine (Sak-
agami and Hayashida 1961).
The nests of L. (E.) matianense were not
randomly distributed in the banks next to
the livestock trails. The great majority of
nests were in the topmost layer of soil at all
of the 31 sites examined. Nests were made
in a 10-15 cm thick zone of dark, loamy,
moist soil, among the tough, fibrous rootlets
of the short turf grasses and forbs. The en-
trances to many nests were hidden, and
partly shaded by an overhanging thatch of
grass blades and dangling rootlets (Fig. 3).
At site 3, where nests were the densest
(2403 per m7), nest entrances were as close
as 2 mm. This made orientation difficult for
returning foragers, which hovered, zigzag-
ging in front of the bank, before finding
their nests. About 150 flying, humming,
bees per m? were visible at a glance. This
is one of the higher nest densities among
bees. Another solitary species, L. (Evy-
laeus) sp. nr. fulvicorne (Kirby) nests in ag-
gregations that reach 101 nests per m? (xX =
54 per m’; Maeta 1966).
In order to study the reason for the lim-
ited dispersal of the nests of L. (E.) mati-
anense, two calibrated, bimetallic, dial
probe thermometers were inserted at 2 lo-
cations into the soil at site 3 and kept there
all day. One thermometer was inserted to
depths of 5, then 15 cm, into the moist,
loamy rootlet zone where the maximum
nests had been made. The other thermom-
eter was inserted near the first, at 5, then 15
cm, in a drier, sandy area about 30 cm be-
low the first, where there were no nests.
Temperatures were recorded periodically at
5 and 15 cm in the nest zone and at the
same times in the nestless zone. The moist,
loamy soil that was selected by the bees for
nesting maintained a more constant and
lower daily temperature than the sandy soil
that was avoided by the bees (Fig. 4). Ev-
idently, the insulating thatch of grass and
the moisture retained by the rootlets buf-
fered the nests and brood cells (mean depth,
6.2 cm) from daily temperature extremes
and desiccation.
167
NEST ARCHITECTURE
Fourteen complete, open nests with
brood cells and 23 miscellaneous cells of L.
(E.) matianense were excavated (Fig. 3).
The circular nest entrances were 2.5 to 3.0
mm in diameter, and were somewhat irreg-
ular, without any noticeable modifications
made by their inhabitants. The main tunnels
were sinuous, 4.0—4.5 mm in diameter, and
2.5 to 11.0 (kX = 6.2) cm deep, ending in a
cell. Eleven nests terminated at one cell,
and 3 nests had 2 cells each. The nest tun-
nels were intertwined among each other and
among rootlets and stones. The friable, fine,
loose soil and rootlets made it difficult to
trace individual tunnels.
The contents of undamaged brood cells
varied. Five cells were new, open, and un-
provisioned, smelling of the characteristi-
cally tangy Dufour’s gland secretion, which
is used by the bees to waterproof the inte-
rior of cells. Marks made by pygidial plates
could be seen in their shiny, smooth inte-
riors. The slightly milky, transparent se-
creted cell linings penetrated the surround-
ing soil, which contained flat mica particles
that formed a terrazzo-like pattern in cells.
The cells were of the usual halictine shape
(Fig. 3). Five cells were open, containing
small, incomplete balls of moist, medium-
yellow apple pollen, with a dusting of
loose, dry pollen. Ten cells were sealed
with a plug of loose soil. Each of these con-
tained a moist, medium-yellow flattened
spheroid of pollen with a groove on top
containing an arched, white 2.0 mm-long
egg. These pollen balls were 3.5—4.0 mm
in diameter and 2.0—2.5 mm high. They
contained 95% apple pollen (n = 5). Brood
cells were 7.5-9.0 mm long, and 4.0—4.5
mm in maximum width, with a 2.0—2.5 mm
neck. No cells contained larvae, pupae, or
adults. The distal ends of about 80% of
cells extended into the bank, but the ends
of some cells were directed toward the front
of the bank, when burrowing bees had en-
countered rocks. Some cells were built
168 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ae
6
Fig. 3. Nests of L. (E.) matianense among the fibrous rootlets of grasses in the loamy, moist, upper 10—15
cm of a sunlit bank next to a livestock trail.
against rocks, separated from them by 1
mm of soil.
There was no evidence of social behav-
ior, which is unlikely to develop, due to the
brief growing season. One nest contained a
female cleptoparasitic halictine, Sphecodes
sp.; others were empty or contained a single
L. (E.) matianense female. No male L. (E.)
matianense was collected at the nest site or
in the orchards. There were few traces of
VOLUME 99, NUMBER 1
°
30
169
Fig. 4. Thermoregulation may be accomplished by the placement of nests. The moist, loamy soil in the
upper 20 cm of the bank among the rootlets where L. (E.) matianense nests (N) stays cooler, and temperatures
there, at 5 cm and 15 cm, fluctuate less, than in the lower, drier, and sandier portion where the bees did not nest
(dashed lines). Diurnal air (stippled area) and soil (lines) temperatures were recorded on May 7.
abandoned cells from previous years’ nest-
ing. Frass from one such cell contained ap-
ple pollen exine.
CONCLUSION
The simple, combless nests of L. (E.) ma-
tianense resemble the nests of the Holarctic,
solitary, boreal species L. (E.) borealis
(Svensson et al. 1977; Sakagami and Toda
1986). Other similarly combless, solitary,
boreal species are L. (E.) sakagamii which
ranges from northern Japan to Manchuria
(Sakagami et al. 1982), and L. (E.) allodal-
um Ebmer et Sakagami, from northern Ja-
pan (Sakagami et al. 1985). The other sol-
itary, Palearctic species of Evylaeus make
earthen combs of cells; for example, L. (E.)
nupricola Sakagami, a boreo-alpine, ice-
age relic in northern Japan (Sakagami
1988), and L. (E.) calceatum, which is sol-
itary at high, cold elevations, but is eusocial
in more temperate zones (Sakagami and
Munakata 1972).
Himalayan apple growers could improve
pollination by providing habitat for nests of
L. (E.) matianense near their orchards. The
trees should be grown on, or near, south-
facing, sunny, turf-covered slopes, where
170 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
vertical cuts have been made, to expose
bare soil banks for nesting bees.
ACKNOWLEDGMENTS
I thank my husband, Lekh R. Batra, for
his help with the research, and for essential
logistic support in this remote area. The re-
search was partly funded by the Foreign
Agricultural Service, O.I.C.D., U.S.D.A. I
thank PA.W. Ebmer of Austria for identi-
fying the bees. E.M. Barrows of George-
town University and B.B. Norden of the
Smithsonian Institution improved the
manuscript. In memory of Shdichi EK Sak-
agami.
LITERATURE CITED
Batra, S. W. T. 1966. Nests and social behavior of
halictine bees of India. Indian Journal of Ento-
mology 28: 375-393.
. 1990a. Bionomics of Evylaeus comagenensis
(Knerer and Atwood) a facultatively polygynous,
univoltine, boreal halictine bee. Proceedings of
the Entomological Society of Washington 92:
725-731.
1990b. Bionomics of a vernal solitary bee,
Andrena (Scrapteropsis) alleghaniensis Viereck,
in the Adirondacks of New York. (Hymenoptera:
Andrenidae). Journal of the Kansas Entomological
Society 63: 260-266.
1996. Biology of Apis laboriosa Smith, a
pollinator of apples at high altitude in the Greater
Himalaya of Garhwal, India (Hymenoptera: Api-
dae). Journal of the Kansas Entomological Soci-
ety. 69: 177-181.
. (in press) Fruit-pollinating bees of the Ga-
rhwal Himalaya, U.P., India. Acta Horticulturae.
Boyle, R. M. D. and B. J. R. Philogéne. 1983. The
native pollinators of an apple orchard: variations
and significance. Journal of Horticultural Science
58: 355-363.
Boyle-Makowski, R. M. D. 1987. The importance of
native pollinators in cultivated orchards: Their
abundance and activities in relation to weather
conditions. Proceedings of the Entomological So-
ciety of Ontario 118: 125-141.
Maeta, Y. 1966. Biological studies on the solitary bee,
Lasioglossum sp. near fulvicorne (Kirby), (1.) en-
vironments of the nesting site and annual devel-
opment of the colony. Kontya 34: 223-231.
Mani, M. S. 1962. Introduction to High Altitude En-
tomology. London: Methuen Co. 302 pp.
. 1978. Ecology and Phytogeography of High
Altitude Plants of the Northwest Himalaya. New
Delhi: Oxford and IBH Publ. 205 pp.
Packer, L. and G. Knerer. 1985. Social evolution and
its correlates in bees of the subgenus Evylaeus
(Hymenoptera: Halictidae). Behavioral Ecology
and Sociobiology 17: 143-149.
Polunin, O. and A. Stainton. 1984. Flowers of the
Himalaya. Delhi: Oxford University Press. 580
Pp.
Sakagami, S. EF 1988. Bionomics of the halictine bees
in northern Japan IV. Lasioglossum (Evylaeus) nu-
pricola sp. nov., a climatic relic. Kontya 56: 337—
3)5)3):
Sakagami, S. E, A. W. Ebmer, and T. Matsumura.
1982. Bionomics of the halictine bees in northern
Japan. Il. Lasioglossum (Evylaeus) sakagamii
(Hymenoptera, Apoidea, Halictidae), with taxo-
nomic notes on allied species. Kontyi 50: 198—
Zale
Sakagami, S. EK and K. Hayashida. 1961. Biology of
the primitive social bee, Halictus duplex Dalla
Torre, III. Activities in spring solitary phase. Jour-
nal of the Faculty of Science, Hokkaido Univer-
sity, Series VI, Zoology 14: 639-682.
Sakagami, S. F, T. Matsumura, and Y. Maeta. 1985.
Bionomics of the halictine bees in northern Japan
Ill, Lasioglossum (Evylaeus) allodalum, with re-
marks on the serially arranged cells in the halic-
tine nests. Konty 53: 409-419.
Sakagami, S. EF and M. Munakata. 1972. Distribution
and bionomics of a transpalearctic eusocial hal-
ictine bee, Lasioglossum (Evylaeus) calceatum, in
northern Japan, with reference to its solitary life
cycle at high altitude. Journal of the Faculty of
Science, Hokkaido University, Series VI, Zoology
18: 411-439.
Sakagami, S. EF and M. J. Toda. 1986. Some arctic
and subarctic solitary bees collected at Inuvik and
Tuktoyaktuk, N.W.T., Canada (Hymenoptera:
Apoidea). Canadian Entomologist 118: 395-405.
Scott-Dupree, C. D. and M. L. Winston. 1987. Wild
bee pollinator diversity and abundance in orchard
and uncultivated habitats in the Okanagan Valley,
British Columbia. Canadian Entomologist 119:
735-745.
Svensson, B. G., P. A. W. Ebmer, and S. FE Sakagami.
1977. Lasioglossum (Evylaeus) boreale, a new
Halictinae (Hymenoptera: Apoidea) species found
in northern Sweden and on Hokkaido, Japan, with
notes on its biology. Entomologia Scandinavica 8:
219-229.
Whiteman, P. T. S. 1985. The mountain environment:
an agronomist’s perspective with a case study
from Jumla, Nepal. Mountain Research and De-
velopment 5: 151-162.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 171-173
A NEW SPECIES OF PSEUDONOMONEURA BEQUAERT
(DIPTERA: MYDIDAE) FROM MEXICO
ScottT J. FITZGERALD AND Boris C. KONDRATIEFF
Department of Entomology, Colorado State University, Fort Collins, CO 80523, U.S.A.
Abstract.—A new mydid fly, Pseudonomoneura calderwoodi, n. sp., is described and
illustrated from 18 specimens collected in Baja California, Mexico. This species is most
similar to Pseudonomoneura nelsoni Fitzgerald and Kondratieff, and the characters for
their separation are summarized. The existing taxonomic key to species of Pseudono-
moneura is modified to accomodate the new species.
Key Words:
Fitzgerald and Kondratieff (1995) pro-
vided a recent review of the mydid fly ge-
nus Pseudonomoneura Bequaert in which
six species are recognized. Through the
kindness of J. A. Calderwood, Santa Bar-
bara Museum of Natural History, 18 males
of Pseudonomoneura which represented a
new species were made available to us for
description.
Terminology of male terminalia follows
Fitzgerald and Kondratieff (1995). Abbre-
viations for depositories of specimens are:
California Academy of Sciences, San Fran-
cisco (CAS); C. P. Gillette Museum of Ar-
thropod Diversity, Colorado State Univer-
sity, Fort Collins (CSU); J. A. Calderwood
Collection, Santa Barbara, California
(JAC); Santa Barbara Museum of Natural
History, Santa Barbara, California (SBMN).
Pseudonomoneura calderwoodi
Fitzgerald and Kondratieff,
New Species
(Figs. 1-3)
Types.—Holotype: 3 (CAS), Mexico:
Baja California [Baja California Norte], 13
km North of Guerrero Negro, dunes, 13
April 1995, J. A. Calderwood. Paratypes:
Mydidae, Pseudonomoneura, Mexico
same data as holotype, 5 ¢ (CAS), 5 6
(CSU), 3 6 (JAC), 4 d (SBMN).
Description.—Male: Head: Frons dense-
ly silver-white tomentose and pilose. Occi-
put black. Antenna short, stout, brown to
black with sparse white pile on pedicel and
scape. Mouthparts vestigial. Thorax: Meso-
notum densely silver-white tomentose with
three opaque gray-brown tomentose vittae.
Lateral vittae narrower and shorter than me-
dial vitta. Mesonotum densely silver-white
pilose except on vittae. Pleurae shining
brown with some areas of silver-white to-
mentum. Halter yellow-white. Scutellum
silver-white tomentose. Legs: Light brown
with femora darker than tibiae and tarsi.
Hind leg with light brown hair and weakly
developed flexor spines. Hind femur not
swollen. Wing: Hyaline, 5—6.5 mm, vena-
tion as in other Pseudonomoneura. Abdo-
men: Tergites brown in ground color with
yellow-white posterior margins. All tergites
with thin silver-white tomentum and de-
cumbent pile except on a middorsal longi-
tudinal brown vitta which runs length of ab-
domen. Sternites brown with sparse decum-
bent pile. Bullae small, dark brown. Geni-
talia: In dorsal view, upper forceps of
epandrium slender, apically rounded (Fig.
172 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-3.
1), in lateral view, apically truncate, simple
(Fig. 2). Gonocoxites slightly cleft with
ventral processes absent (Fig. 3).
Female: Unknown.
Etymology.—The patronym honors Mr.
J. A. Calderwood (SBMN), who collected
and made available specimens for descrip-
tion.
Diagnosis.—The vestigial mouthparts
Pseudonomoneura calderwoodi, male terminalia. 1, Dorsal view. 2, Lateral view. 3, Ventral view.
and the simple form of the upper forceps of
the epandrium (i.e. lacking the digitate
structure of P. californica (Hardy), the bi-
furcate structure of P. bajaensis Fitzgerald
and Kondratieff, or the upper and lower
flanges of P. hirta (Coquillett), P. miche-
neri (James) and P. tinkhami (Hardy)) will
easily separate males of P. calderwoodi
from all other Pseudonomoneura, with the
VOLUME 99, NUMBER I
exception of P. nelsoni Fitzgerald and Kon-
dratieff. Males of P. calderwoodi can be
distinguished from the similar P. nelsoni,
by the genitalia in dorsal view which have
the upper forceps of the epandrium apically
rounded (Fig. 1), rather than apically acute
and bent outward (see Fitzgerald and Kon-
dratieff 1995: 32, Fig. 21). In lateral view,
the upper forceps of the epandrium are
more apically truncate (Fig. 2) than in P.
nelsoni (see Fitzgerald and Kondratieff
19952 32; Fig: 20):
Males of P. calderwoodi can be identified
using the key of Fitzgerald and Kondratieff
(1995) with the following modification of
couplets | and 2 (figure numbers are those
in Fitzgerald and Kondratieff 1995):
[eee Vlouthpartsivesticial) = 244.65 see eee 2
— Mouthparts well-developed
Ms
2. Male terminalia with upper forceps of epan-
driumssimplei(Gigsy20) 21), 22.) te ele aes 2a
— Male terminalia with upper forceps of epan-
drium bifurcate (Figs. 17, 18) ....... bajaensis
2a. Upper forceps of epandrium apically acute
and bent outward in dorsal view (Fig. 21)
FE ee EMR ft AH AEA) ee MS) EEE sia nelsoni
— Upper forceps of epandrium apically rounded
and not bent outward in dorsal view ........
ACKNOWLEDGMENTS
We thank Mr. J. A. Calderwood for mak-
ing material available for study. Lynn C.
Bjork provided the illustrations.
LITERATURE CITED
Fitzgerald, S. J. and B. C. Kondratieff. 1995. A re-
view of the mydid genus Pseudonomoneura Be-
quaert (Diptera: Mydidae), with the description of
two new species. Proceedings of the Entomolo-
gical Society of Washington 97: 22—34.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 174-179
REPORT ON A COLLECTION OF BETHYLIDAE (HYMENOPTERA) FROM
CENTRAL FLORIDA
HOwARD E. EVANS AND STUART M. FULLERTON
(HEE) Department of Entomology, Colorado State University, Fort Collins, CO 80523,
U.S.A.; (SMF) Department of Biology, University of Central Florida, Orlando, FL 32816-
23508, WES.
Abstract.—About 3200 specimens representing 51 species of Bethylidae were taken,
primarily in Malaise traps, at 5 sites in Orange Co., and one site in Seminole Co., Florida.
These are listed, with habitats and dates of collection. One new species is described:
Bakeriella mira Evans. Range extensions are indicated for four species, and one new
synonymy is made.
Key Words:
Beginning in 1990, biologists at the Uni-
versity of Central Florida (UCF) have sur-
veyed the arthropod fauna of the campus,
using Malaise traps and a variety of other
sampling techniques. The university was
founded in 1963 on an 1100 acre plot of
land that had previously been a cattle ranch.
Malaise traps were set up in four ecologi-
cally distinct sites on campus and were
maintained throughout the year, being emp-
tied every 4-5 days. Specimens were col-
lected in cyanide prior to 1995, in that year
and 1996 in 70% isopropyl alcohol. Traps
were of the design of Townes (1972) with
some modifications. Fine-meshed cloth was
used, in some cases all black, in others
black and white. Two Malaise traps were
also set up outside the campus, one at Rock
Springs Run State Reserve, 40 km NW of
the UCF campus, and one at the home of
SMF in Oviedo, 9 km N of the campus. The
last locality is in Seminole Co., the others
in Orange Co.
A report on the Bethylidae seems justi-
fied because of the large number of speci-
mens taken (3200) and the diversity of the
fauna (51 species). Some species occurred
Hymenoptera, Bethylidae, wasps, Florida
in remarkable numbers, for example 817
males and 9 females of Pseudisobrachium
flaviventre (Kieffer). Such figures suggest
that some Bethylidae are much more plen-
tiful than commonly realized. To the best of
our knowledge, no species of Pseudiso-
brachium has ever been reared from a host.
Females have been found in ant nests, but
it is uncertain whether they attack the lar-
vae of ants or of their beetle myrmeco-
philes. Clearly there is much room for re-
search.
The collection includes one previously
undescribed species and four range exten-
sions, two of them unexpected reports of
species of more tropical distribution. One
new synonymy is made on the basis of se-
ries of males and females formerly assigned
to different species.
COLLECTION SITES
The collection sites are referred to by
number in the list that follows.
1. A disturbed area of long leaf and slash
pine (Pinus palustris Mill. and P. elliottii
Engelm.), turkey oak (Quercus laevis Wal-
ter), and saw palmetto (Serenoa repens
VOLUME 99, NUMBER 1
(Bartr.). This site has not been burned for
many years and now bears many sand
pines, Pinus clausa (Engelm.) Sarg. UCF
campus.
2. A scrub growing on fossil sand dunes
with vegetation that includes sand pine,
Florida rosemary (Ceratiola ericoides
Michx.), saw palmetto, and scrub oaks
(Quercus geminata Small, Q. myrtifolia
Willd., and Q. chapmani Sargent, in order
of abundance), and lyonias, Lyonia ferru-
ginea (Walt.) Nutt. and L. lucida (Lam.)
Koch. UCF campus.
3. Cypress dome, consisting of pond cy-
press (Taxodium ascendens Brogn.), mai-
dencane (Panicum hemitomon Schultes),
wax myrtle (Myrica cerifera L.), dahoon
holly Ulex cassine L.), and gallberry (Ilex
glabra L.). The trap was about 3 m out in
the water at the middle of the cypress ring.
UCF campus.
4. Site near a nyssa pond (Nyssa sylva-
tica Marsh) that was wet for a large part of
the year; an area adjacent to the site has
since been developed as a retention pond.
Vegetation consists of long leaf and slash
pine, saw palmetto, shining lyonia, Lyonia
lucida (Lam.) Koch, and assorted grasses.
UCF campus.
5. Rock Springs Run State Reserve. This
is a 356 acre parcel of land that has not
been burned within the last 40-50 years.
Predominant vegetation is sand pine and
sand live oak (Quercus geminata Small),
with a secondary mixture of myrtle oak (Q.
myrtifolia Willd.), Chapman’s oak (Q.
chapmani Sargent), saw palmetto, and Flor-
ida rosemary.
6. A rural yard in an older neighborhood
of Oviedo. One side of the trap faced a
moderately cultivated yard, with weeds and
a mixture of native and exotic plants; the
other side faced an abandoned field that was
once a citrus orchard.
LIST OF SPECIES
Listing of genera follows Evans (1978),
with species of each genus listed alphabet-
ically.
175
SUBFAMILY PRISTOCERINAE
Pristocera armifera (Say). 7 22, 87 36.
May-—Dec. Sites 1, 2, 4.
P. atra Klug. 2 22, 218 36. Throughout
the year, Sites 15 2; 5:
P. bridwelli Evans. 1 6. June. Site 1. New
record for Florida.
P.fraterna Evans. 9) 2 or
Throughout the year, all sites.
Apenesia parapolita (Evans). 2 2°, 30
66. Throughout the year. Sites 1, 2, 5.
Dissomphalus apertus Kieffer. 21 46.
May-Sept. Sites 3, 4.
D. barberi Evans. 1 3. Oct. Site 6. New
record for Florida.
D. kansanus Evans. 1 ¢. Oct. Site 1.
Pseudisobrachium arenarium Evans. 807
36. Throughout the year. Sites 1, 2, 5,
6. See note below.
P. ashmeadi Evans. 73 36. Jan.—Feb.,
June—Dec: Sites 15 2, 4, 5;.6:
P. carolinianum Evans. 59 3 6. Jan., June—
Nov: Site 5:
P. flaviventre (Kieffer). 9 22, 817 oo.
Throughout the year. Sites 1, 2, 4, 6. See
note below.
P. rufiventre (Ashmead). 49 o<¢. Jan.,
Apr.—Dec. Sites 1, 2, 5, 6.
Wa 6 O.
SUBFAMILY EPYRINAE
Rhabdepyris amabilis Fouts. 5 ¢¢. May—
Aug. Sites 2, 4, 5.
R< carohnianus, Evans, 8 22, 35: did:
May-Sept. Sites 1, 2.
R. mellipes Evans. 1 2. May. Site 4.
R. muesebecki Evans. 3 366. June, July.
Sites 1,3:
Anisepyris analis (Cresson). 59 22, 29
36. Jan.—Nov. Sites 1, 2, 4, 5, 6.
A. bradleyi (Evans). 1 @. July. Site 2.
A. columbianus (Ashmead). 6 2 2, 18 36.
Jan., Apr.—Aug. Sites 1, 2, 5.
A. gibbosifrons Evans. 1 ¢. June. Site 6.
Flight intercept trap.
A. grandis (Ashmead). 31 22, 210 dd.
Throughout the year. All six sites.
A. subviolaceus Kieffer. 1 2, 8 ¢¢. May-—
Aug. Sites: 1;)2, 4, 5:
176 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Epyris alachua Evans. 4 2? ¢. Mar., May,
July. Sites 22 6:
E. deficiens Krombein. 4 2¢@, 20 dodo.
Throughout the year. Sites 1, 2, 5.
E. festivus Evans. 29 2 2, 24 66. Through-
Outathesyeat. Sites 1, 2, 3, 5: See mote
below.
E. myrmecophilus (Brues). 1 2, 13 66.
Throughout the year. Sites 1, 2.
E. oriplanus Kieffer. 1 2, 2 6d. May, July,
Deca sitessl; 2.
E. rufipes (Say). 39 2 2, 182 66. Through-
out the year. All six sites.
E. spissus Evans. 20 °°, 6 66. Through-
out the year Sites 1, 2, 3.
E. tricostatus Evans. 1 2, 1 ¢. July, Aug.
Sites 15 2.
Bakeriella mira Evans, n. sp. described be-
low. 2 366. June, July. Site 1.
Holepyris floridanus (Ashmead). 3 2 2, 18
36. Throughout the year. Sites 1, 2, 5, 6.
H. graminis Evans. 1 2,7 66. Jan., June—
Nove Sites. 1, 2, 4, .5,.6.
H. hispaniolae Evans. 3 22, 1 3d. Apr—
July. Sites 1, 2,5. Two 2 @ taken in flight
intercept trap. See note below.
H. lautus Evans. 1 2, 21 66. Throughout
the year. Sites 1, 2, 4, 5, 6.
H. micidus Evans. 2 22,9 3d. May, July—
Dec sites |. 2,,6:
H. subapterus (Melander and Brues). 2 2 2.
May. Site 2.
H. subtilis Evans. 1 2, 3 dd. May, Aug.,
Sept. Sites 1, 2, 3. 2 taken in flight in-
tercept trap. See note below.
Laelius centratus (Say). 1 2. April. Site 6.
Scleroderma macrogaster (Ashmead). 3
22. Apr.—May. Site 6.
SUBFAMILY BETHYLINAE
Prosierola bicarinata (Brues). 1 2. July.
Site 6.
Goniozus columbianus Ashmead. 1 @. Feb.
Site 4.
G. floridanus (Ashmead). 1 2. Aug. Site 2.
G. fratellus Evans. 2 2 2. Mar., Oct. Site 2.
G. hortorum Brues. 3 2 2. Mar., May, Aug.
Sites 1, 4.
G. hubbardi Howard. 6 2. Feb., Mar.,
June; "DecrSites 15°45 16:
G. indigens Evans. 5 2&2. May. Site 2.
Flight intercept trap.
G. nigrifemur Ashmead. 10 @ @. Feb., Mar.,
May siSept Oct'Sites) 1921356:
G. scitulus Evans. 3 22. Apr., Aug., Oct.
Sites 2, 6.
G. seminole Evans. 9 2 2. Jan., Feb., Apr.,
June—Nov. Sites 2, 4, 5.
NOTES AND DESCRIPTIONS OF
INDIVIDUAL SPECIES
Pseudisobrachium arenarium Evans
This species is a member of the prolon-
gatum group of Evans (1961), in which the
mandibles of the male are 5-toothed, but the
third and fourth teeth are very small and
may be partially or even wholly fused, re-
sulting in 4-toothed mandibles. Of the 807
males collected in this study, 233 (29%)
have only four distinct mandibular teeth.
These are mainly smaller males and are eas-
ily confused with rufiventre. However, in
arenarium the minimum width of the frons
barely if at all exceeds eye height and the
antennae are more elongate, segment 3 be-
ing about twice as long as wide.
Pseudisobrachium flaviventre (Kieffer)
Correct association of the sexes in this
genus is difficult, as males are commonly
taken sweeping, in traps, or at light, while
females are usually found in ant nests. In
this case the 9 females may have been car-
ried into malaise traps by males through
phoretic copulation, as is known to occur in
other Pristocerinae. These females are min-
ute, varying in body length from 2 to 3 mm,
in head length from 0.40 to 0.53 mm;
length of the head varies from 1.25 to 1.42
times width of the head. This stands in con-
trast to the female rufiventre, in which the
head is 0.68 to 0.8 mm long and 1.1 to 1.2
times as long as wide. The mandibles of the
9 flaviventre females are basically biden-
tate, but in 5 specimens a small knob basad
of the uppermost tooth can be detected, and
VOLUME 99, NUMBER 1
in 2 a small third tooth is present. By and
large, these specimens key well to flaviven-
tre in the keys to females presented by
Evans (1961, 1978).
Some of the characters previously used
for identification of males have proved of
doubtful reliability. The most useful feature
for identifying male flaviventre is the com-
pact, pale yellowish to reddish-brown an-
tennae, in which the outer flagellar seg-
ments (except the last) are no longer than
wide. In contrast, male rufiventre have
brown antennae in which the outer flagellar
segments are distinctly longer than wide.
Epyris festivus Evans
This is a member of the depressigaster
group of Evans (1969), in which the ab-
domen of the female is strongly depressed
and bears one or more hyaline plates ven-
trally. In characterizing the group, Evans
suggested the possibility that members of
the idionotum group might represent the
male sex. Collection of 29 females of fes-
tivus and 24 males of idionotum at similar
times and sites makes it clear that these are
female and male of one species. While
pronotal structure is very different, there is
similarity in the dark olive-green coloration
as well as in the 9 or 10 parallel propodeal
carinae. This is a new synonymy, festivus
having page priority over idionotum; both
were described by Evans in 1969.
Bakeriella mira Evans, new species
Holotype.—d, FLORIDA: Orange Co.,
UCE Orlando, VII-22-93; LLP-sand pine,
turkey oak; Malaise trap, S.M. Fullerton
collector (National Museum of Natural His-
tory, Smithsonian Institution). Paratype.—
3d, same data except collected VI-7-96
(Florida State Collection of Arthropods).
Description of holotype male.—Length
4.2 mm, fore wing 2.7 mm. Black; antenna
bright reddish-brown, apical two segments
darkened; palpi light brown; mandible
black basally, reddish apically; tegula light
brown; coxae and mid and hind femora
black, fore femur and all tibiae and tarsi
177
brown; wings hyaline. Median lobe of clyp-
eus obtusely angulate; antennal scrobes not
carinate; genae not carinate. Scape about
twice as long as thick; first 4 antennal seg-
ments in a ratio of 17:7:7:10. Head about
as wide as high; eyes weakly convergent
below, not hairy; width of frons 0.58 times
width of head, 1.10 times height of eye.
Ocelli in a right triangle, ocello-ocular line
subequal to width of ocellar triangle. Frons
weakly shining, alutaceous, with small, se-
tigerous punctures separated by somewhat
more than their own diameters.
Thoracic dorsum similarly alutaceous
and punctate, bearing short, brown hairs.
Pronotum with anterior lateral angles sharp
and slightly protuberant; lateral margins
concave, paralleled internally by a foveo-
late groove; anterior margin of disc elevat-
ed, the elevation extended broadly back-
ward, then slightly emarginate, margins of
the elevation foveolate; posterior pronotal
margin simple (Fig. 1). Mesoscutum with
deeply impressed, somewhat foveolate no-
tauli that are progressively more slender an-
teriorly, reaching the pronotum as _ thin
lines. Scutellum flat, with two large pits
separated by slightly less than their own di-
ameters. Propodeal disc 1.15 times as wide
as long, with a complete median carina
flanked by 4 incomplete, more irregular ca-
rinae between which the surface is foveo-
late; sides of disc weakly transversely stri-
ate, posterior margin carinate. Mesopleu-
rum with a large, bowl-shaped median pit
and two smaller anterior pits; lower margin
with a weakly foveolate longitudinal carina
(ies 2):
Variation.—The paratype is slightly larg-
er than the type, body length measuring 4.9
mm, fore wing 2.9 mm. Resemblance to the
type is very close, but there are two minor
differences in sculpturing: the scutellar pits
are not quite as wide and are separated by
approximately their greatest diameters; and
there are only 3 well-defined propodeal ca-
rinae, the 2 irregular carinae close beside
the median carina being less well formed
than in the type.
178
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Bisse ley?
Bakeriella mira, holotype. 1, Dorsal aspect of thorax and propodeum. 2, Lateral aspect of me-
sopleurum, anterior margin to the right. Figures not drawn to same scale.
Remarks.—This striking species is
known from only two specimens. The form
of the anterior elevation of the pronotum is
unique in the genus. Well separated scutel-
lar pits such as those in mira occur in only
one other known species of Bakeriella, cris-
tata Evans, known from Brazil, Bolivia,
and Argentina (Evans, 1964, 1979). Only
one other species of Bakeriella is known
from the United States: floridana Evans, re-
ported from Dade Co., Florida, also from
Jamaica. This species differs in many de-
tails from mira: the scutellar pits are sepa-
rated by a thin septum, the pronotum has a
transverse carina across the front, and the
mesopleurum lacks a large median depres-
sion.
Twenty-three species of this neotropical
genus are now known. Evans (1979) pro-
vided a key for 18 of these. Azevedo (1991,
1994) has recently added three species of
Bakeriella from Brazil and presented a key
to the 11 species having an anterior trans-
verse carina on the pronotum.
Holepyris hispaniolae Evans
This species was described by Evans
(1977) from a single female from Constan-
za, Dominican Republic. Three females
from sites 2 and 5 agree very closely with
the type and represent a notable range ex-
tension—not however without precedent,
since Bakeriella floridana Evans and Ani-
sepyris aurichalceus (Westwood) also occur
both in Florida and in the West Indies. A
male from site | almost certainly represents
the previously unknown male of this spe-
cies. In the key of Evans (1958) it runs to
floridanus (Ashmead) but the prominently
5-carinate propodeum (similar to that of the
female) and the broader head distinguish it
readily. The head is about as wide as high;
width of the frons is 0.61 times width of
the head, 1.27 times height of the eye. The
VOLUME 99, NUMBER 1
antennae are wholly brown, the coxae
black, femora light brown, tibiae and tarsi
testaceous.
Holepyris subtilis Evans
This species was described from Costa
Rica and Panama, with a single female
from southern California and a single male
from southern Arizona (Evans, 1978). It is
a distinctive species, males lacking erect
pubescence on the flagellum and having 5
propodeal carinae with two shorter carinae
laterad of these. Females are one of only
two American species lacking a transverse
pronotal carina; the frons is unusually nar-
row, the minimum width in this instance
only 0.70 times the eye height; the antennal
scape is fully 6.5 times as long as thick.
Three males from sites | and 3 and a female
from site 2 represent a major range exten-
sion for this species.
ACKNOWLEDGMENTS
We thank Karl V. Krombein, Smithsoni-
an Institution, for lending us several types
from the National Museum of Natural His-
tory. We also thank Jon C. Longhurst and
David W. Hanf for help with the Malaise
trap monitoring.
179
LITERATURE CITED
Azevedo, C. O.
Bakeriella Kieffer (Hymenoptera, Bethylidae) da
regiao de Sao Carlos, SP. Revista Brasileira de
Entomologia 35: 535-538.
1994. Descriptions of two new species and
notes on the genus Bakeriella Kieffer from Brazil
and Ecuador (Hymenoptera: Bethylidae). Journal
of Hymenoptera Research 3: 145—150.
Evans, H. E. 1961. A revision of the genus Pseudi-
sobrachium in North and Central America (Hy-
menoptera, Bethylidae). Bulletin of the Museum
of Comparative Zoology Harvard 125: 211-318.
. 1964. A synopsis of the American Bethylidae
(Hymenoptera, Aculeata). Bulletin of the Museum
of Comparative Zoology Harvard 132: 1—222.
. 1969. A revision of the genus Epyris in the
Americas (Hymenoptera: Bethylidae). Transac-
tions of the American Entomological Society 95:
181-352.
1977. A revision of the genus Holepyris in
the Americas (Hymenoptera: Bethylidae). Trans-
actions of the American Entomological Society
103: 531-579.
1978. The Bethylidae of America north of
Mexico. American Entomological Institute Mem-
oir 27, 332 pp.
1979. A reconsideration of the genus Bak-
eriella (Hymenoptera: Bethylidae). Journal of the
New York Entomological Society 87: 256—266.
Townes, H. 1972. A light-weight Malaise trap. Ento-
mological News 83: 239-247.
1991. Espécie nova e notas sobre
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 180-193
LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF
PROCECIDOCHARES ANTHRACINA (DOANE) (DIPTERA: TEPHRITIDAE)
ON SOLIDAGO CALIFORNICA NUTTALL IN SOUTHERN CALIFORNIA
RICHARD D. GOEDEN AND JEFFREY A. TEERINK
Department of Entomology, University of California, Riverside, CA 92521, U.S.A.
Abstract.—Procecidochares anthracina (Doane) is a univoltine, circumnatal, tephritid
fly widely distributed in the western United States and apparently nearly monophagous
on Solidago californica Nuttall and as yet unknown congeners probably of similar growth
habit. Unique among other Procecidochares species studied to date, P. anthracina repro-
duces gregariously in underground bud galls on shoots arising from host-plant rhizomes.
The egg, first through third instars, and puparium are described and figured for the first
time. The small, rounded, rugose pads that circumscribe the prothorax; the presence of
four, small subdorsal sensilla ventrolaterad of the dorsal sensory organ and dorsal to the
anterior sensory lobe, and the lateral spiracular complexes distinguish the third instar from
other gallicolous Procecidochares examined to date. Pupariation occurs within subsurface
galls and teneral adults must climb upward through several cm of humus to free them-
selves and mate. Premating, mating, and postmating behaviors are described which include
spurts of exceptionally rapid, blurred wing enantions. Mated females must reenter the
humus layer to oviposit in buds on buried shoots, as subsequently must parasitoids to
locate and probably enter galls to oviposit in the larvae or puparia. Eurytoma obtusiventrus
Gahan (Hymenoptera: Eurytomidae) is reported as a solitary, larval-pupal endoparasitoid
reared from puparia; Pronotalia carlinarium Gradwell (Hymenoptera: Eulophidae) is a
gregarious endoparasitoid of puparia.
Key Words: Insecta, Procecidochares, Solidago, biology, galls, taxonomy of immature
stages, reproductive behavior, parasitoids
Life histories and immature stages of MATERIALS AND METHODS
four species of Procecidochares (Diptera:
Tephritidae) in southern California have
been described to date, i.e. P. flavipes Al-
drich (Goeden et al. 1994a), P. kristineae
Goeden (Silverman and Goeden 1980, Goe-
den and Teerink 1997), P. lisae Goeden
(Goeden and Teerink 1997), and P. stonei
Blanc and Foote (Green et al. 1993). Herein
we describe the life history and immature
Our field studies on P. anthracina were
conducted principally at two locations in
southern California during 1992-1995: (1)
at the junction of Kitchen Creek Road and
the Sunrise Highway at 1720-m elevation,
Cleveland National Forest, San Diego Co.
and (2) in Spilman Canyon SE of Lake
Hemet at 1380 m, San Bernardino National
Forest (S section), Riverside Co. Infested
Stages of a fifth species, P. anthracina (Do-
ane), which uniquely forms bud galls on
shoots arising from subsurface rhizomes of
Solidago californica Nuttall (Asteraceae).
rootstocks dug from humus beneath decid-
uous Oaks (Quercus spp.) within patches of
shoots of Solidago californica were trans-
ported in cold-chests in an air-conditioned
VOLUME 99, NUMBER 1
vehicle to the laboratory and stored under
refrigeration for subsequent dissection, pho-
tography, description, and measurement.
Twenty six eggs dissected from buds and
seven first-, 12 second-, and 14 third-instar
larvae, and six puparia from galls were pre-
served in 70% EtOH for scanning electron
microscopy (SEM). Additional puparia
were placed in separate, glass shell vials
stoppered with absorbant cotton and held in
humidity chambers at room temperature for
adult and parasitoid emergence. Specimens
for SEM were hydrated to distilled water in
a decreasing series of acidulated EtOH.
They were osmicated for 24 h, dehydrated
through an increasing series of acidulated
EtOH and two, 1-h immersions in Hexa-
methlydisilazane (HMDS), mounted on
stubs, sputter-coated with a gold-palladium
alloy, and studied with a JEOL JSM C-35
SEM in the Department of Nematology,
University of California, Riverside.
Most adults reared from isolated puparia
obtained from galls were individually caged
with excavated rhizome fragments attached
or unattached to basal portions of aerial
shoots in 850-ml, clear-plastic, screened-top
cages with a cotton wick and basal water
reservoir and provisioned with a strip of pa-
per toweling impregnated with yeast hy-
drolyzate and sucrose. Those cagings with-
out rhizome bouquets were used for lon-
gevity studies and those with bouquets for
oviposition tests in the insectary of the De-
partment of Entomology, University of Cal-
ifornia, Riverside, at 25 + 1°C, and 14/10
(L/D) photoperiod. Virgin male and female
flies obtained from emergence vials were
paired in clear-plastic petri dishes provi-
sioned with a flattened, water-moistened
pad of absorbant cotton spotted with honey
(Headrick and Goeden 1991) for direct ob-
servations, videorecording, and _still-pho-
tography of their courtship and copulation
behavior.
Plant names used in this paper follow
Munz (1974); tephritid names and nomen-
clature follow Foote et al. (1993). Termi-
nology and telegraphic format used to de-
181
scribe the immature stages follow Goeden
et al. (1993; 1994a, b; 1995a, b) and Goe-
den and Teerink (1996a, b, c; 1997) and our
earlier works cited therein. Means + SE are
used throughout this paper. Voucher speci-
mens of P. anthracina and its parasitoids
reside in the research collections of RDG;
preserved specimens of eggs, larvae and
puparia are stored in a separate collection
of immature Tephritidae maintained by JAT.
RESULTS AND DISCUSSION
TAXONOMY
Adult.—First described by Doane (1899)
as Oedaspis anthracina, then transferred to
Cecidochares by Bezzi (1910), P. anthra-
cina was assigned its current name by Bezzi
and Tavares (1916) (Foote et al. 1993).
Only the wing pattern of the adult has been
figured by Doane (1899), Foote and Blanc
(1963), and Foote et al. (1993). Adults are
separated from other congeners, and ulti-
mately from P. australis Aldrich and P.
grindeliae Aldrich, which this species most
closely resembles, by the characters incor-
porated in the key to North American Pro-
cecidochares in Foote et al. (1993).
Immature stages.—Egg: Twenty eggs of
P. anthracina were white, opaque, smooth;
with an elongate-ellipsoidal body, 0.51 +
0.005 (range, 0.47—0.54) mm long, 0.14 +
0.002 (range, 0.13—0.19) mm wide, smooth-
ly rounded at tapered posterior end (Fig.
1A), and with a peg-like anterior pedicel,
0.02 mm long and a few, large aeropyles
(Fig. 1B). The egg of P. anthracina is sim-
ilar in shape but smaller than P. flavipes
(Goeden et al. 1994a) and P. lisae (Goeden
and Teerink 1997). Procecidochares stonei
(Green et al. 1993) and P. kristineae (Sil-
verman and Goeden 1980, Goeden and
Teerink 1997) eggs are smaller than P. an-
thracina eggs and have a tapered pedicel
with much smaller, oval aeropyles.
Third instar: Third instar of P. anthra-
cina superficially smooth, elongate cylin-
drical, tapering anteriorly, rounded posteri-
orly (Fig. 2A); gnathocephalon conical,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1SKU A468
Fig. 1. Egg of Procecidochares anthracina. (A) habitus; (B) anterior end, aeropyles.
VOLUME 99, NUMBER 1
smooth, lacking rugose pads (Fig. 2B);
paired dorsal sensory organs consist of a
dome-shaped papilla (Fig. 2C-1); four,
small, dome-shaped sensilla, each with a
central papilla, located ventrolaterad of dor-
sal sensory organ (Fig. 2C-2); anterior sen-
sory lobe bears the terminal sensory organ
(Fig. 2C-3), pit sensory organ (Fig. 2C-4),
lateral sensory organ (Fig. 2C-5) and su-
pralateral sensory organ (Fig. 2C-6); stomal
sense organs distinct, composed of raised
lobe with two pore sensilla, ventrad of an-
terior sensory lobes (Fig. 2D-1); lateral sen-
silla laterad of stomal sense organ (Fig.
2D-2), two of eight verruciform sensilla, in-
cluding ventrolateral sensilla, circumscrib-
ing gnathocephalon (Fig. 2B-1); mouth
hooks tridentate, teeth conical (Fig. 2B-2,
D-3); median oral lobe obscured in pre-
pared specimens; prothorax covered ante-
riorly with small, rounded rugose pads (Fig.
2B-3); verruciform sensilla posterior to ru-
gose pads on dorsal half of prothorax (Fig.
2B-4); anterior spiracles on posterior mar-
gin of prothorax consist of two dome-
shaped papillae (Fig. 2E); meso- and me-
tathoracic lateral spiracular complexes con-
sist of an open spiracle and two verruciform
sensilla; abdominal lateral spiracular com-
plexes with an open spiracle (Fig. 2F-1) and
one verruciform sensillum (Fig. 2F-2); cau-
dal segment smooth (Fig. 2G-1); posterior
spiracular plates with three ovoid rimae, ca.
0.03 mm in length (Fig. 2H-1), and four
spiniform interspiracular processes, longest
measuring 0.007 mm (Fig. 2H-2); com-
pound sensilla ventrad of posterior spirac-
ular plates consist of a stelex sensillum
(Fig. 2G-2) and a verruciform sensillum
(Fig. 2G-3).
The P. anthracina third instar differs
from that of the flower head-infesting P.
flavipes in that the mouth hooks are triden-
tate, the rugose pads are limited to the pro-
thorax and no minute acanthae are present
in the intersegmental areas nor on the cau-
dal segment (Goeden et al. 1994a). Proce-
cidochares anthracina more closely resem-
bles the other gall-forming Procecidochar-
183
es Spp. we previously have examined, i.e.,
P. kristineae, P. lisae and P. stonei, but dif-
fers mainly in that its prothorax is circum-
scribed by small, rounded rugose pads;
whereas, the other three gall-forming spe-
cies lack these rugose pads (Goeden and
fcennk 1997-, Green etsaly 1993). Differ
ences also were noted in the lateral spirac-
ular complexes; whereby, P. kristineae and
P. lisae lack verruciform sensilla on the
metathorax (Goeden and Teerink 1997),
and the verruciform sensilla in P. stonei are
anterior to the spiracle (Green et al. 1993).
Several previously undescribed sensilla
were found on the gnathocephalon of P. an-
thracina. Four small sensilla located ven-
trolaterad of the dorsal sensory organ and
dorsad to the anterior sensory lobe are col-
lectively named subdorsal sensilla, and are
unreported among previously examined,
nonfrugivorous Tephritidae (Goeden and
Headrick 1990, 1991a, b, 1992; Goeden et
al. 1993, 1994a, b, 1995a, b; Goeden and
Teerink 19964, b, c; Green et al. 1993:
Headrick and Goeden 1990, 1991, 1993;
Headrick et al. 1995). A single, subdorsal
sensillum located laterad to the dorsal sen-
sory organ was reported in Trupanea ni-
gricornis and T. bisetosa (Knio et al. 1996)
and Dioxyna picciola (Headrick et al.
1996).
Second instar: Second instar of P. an-
thracina superficially smooth, elongate cy-
lindrical (Fig. 3A); gnathocephalon conical,
smooth, lacking rugose pads (Fig. 3B);
paired dorsal sensory organs each consist of
a dome-shaped papilla (Fig. 3C-1); anterior
sensory lobes (Fig. 3B-1) bear the terminal
sensory organ (Fig. 3C-2), pit sensory or-
gan (Fig. 3C-3), lateral sensory organ (Fig.
3C-4), and supralateral sensory organ (Fig.
3C-5); stomal sense organ ventrad of ante-
rior sensory lobe (Fig. 3B-2); mouth hooks
obscured, apical tooth conical (Fig. 3B-3);
median oral lobe obscured in prepared
specimens; prothorax covered anteriorly
with small, rounded rugose pads; verruci-
form sensilla posterior to rugose pads on
dorsal half of prothorax; anterior spiracles
184 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Third instar of Procecidochares anthracina. (A) habitus, anterior to left; (B) gnathocephalon, anterior
view, 1—verruciform sensilla, 2—mouth hooks, 3—rugose pads, 4—prothoracic verruciform sensilla; (C) an-
terior sensory lobe, 1—dorsal sensory organ, 2—subdorsal sensilla, 3—terminal sensory organ, 4—pit sensory
organ, 5—lateral sensory organ, 6—supralateral sensory organ; (D) gnathocephalon, anterior view, 1—stomal
sense organ, 2—lateral sensillum, 3—mouth hooks; (E) anterior spiracles; (F) fourth abdominal lateral spiracular
VOLUME 99, NUMBER 1
each consist of two dome-shaped papillae;
lateral spiracular complexes not observed;
caudal segment circumscribed by stelex
sensilla (Fig. 3D-1); posterior spiracular
plates bear three ovoid rimae, ca. 0.008 mm
in length (Fig. 3D-2), and four spiniform
interspiracular processes, longest measuring
0.005 mm (Fig. 3D-3); compound sensilla
ventrad of spiracular plates consist of a ste-
lex sensillum (Fig. 3D-4) and a verruciform
sensillum (Fig. 3D-5).
Second instars of P. anthracina possess
the same sensory structures as the mature
larva, but the structures are not as well-de-
fined. The mouth hooks, median oral lobe,
and lateral spiracular complexes were ob-
scured in prepared specimens.
First instar: First instars of P. anthracina
cylindrical, rounded posteriorly, minute
acanthae circumscribe segmental lines (Fig.
4A); gnathocephalon conical, smooth (Fig.
4B); dorsal sensory organ consists of dome-
shaped papilla (Fig. 4B-1, C-1); anterior
sensory lobe (Fig. 4B-2) bears terminal sen-
sory organ (Fig. 4C-2), pit sensory organ
(Fig. 4C-3), lateral sensory organ (Fig.
4C-4) and supralateral sensory organ (Fig.
4C-5); stomal sense organs ventrad of an-
terior sensory lobes indistinct (Fig. 4B-3);
mouth hooks obscured in prepared speci-
mens; anterior spiracles absent; lateral spi-
racular complexes not observed; caudal
segment circumscribed by stelex sensilla
(Fig. 4D-1); posterior spiracular plates bear
two ovoid rimae, ca. 0.001 mm in length
(Fig. 4D-2), and four rudimentary interspi-
racular processes, ca. 0.0003 mm in length
(Fig. 4D-3).
The features of the first instar are less
distinct than those of the mature larva. The
first instar lacks the prothoracic rugose
pads, but does have minute acanthae cir-
cumscribing the intersegmental lines not
,
185
found in later instars. The anterior sensory
lobes and dorsal sensory organs comprise
most of the gnathocephalon. The four sub-
dorsal sensilla are not present in the first
instar. As the mouth hooks were obscured
in all prepared specimens, we were unable
to determine if the apical tooth was grooved
as in P. lisae and P. kristineae (Goeden and
Teerink, 1997). The respiratory system of
the first instar was very reduced compared
to both later instars. The posterior spiracu-
lar plates with two small rimae and the in-
terspiracular processes were almost indis-
tinct.
Puparium: Puparium of P. anthracina
elongate ellipsoidal (Fig. 5A), anterior end
bears the invagination scar (Fig. 5A-1), pro-
thoracic rugose pads (Fig. 5B-2), verruci-
form sensilla (Fig. 5B-3) and anterior spir-
acles (Fig. 5B-4); caudal segment smooth,
bears the posterior spiracular plates (Fig.
5C-1), and compound sensilla (Fig. 5C-2).
One hundred and forty five puparia of P.
anthracina averaged 3.56 + 0.03 (range,
2.16—4.68) mm in length; 1.55 + 0.02
(range, 0.94—1.99) mm in width.
DISTRIBUTION AND HOSTS
Because of its unusual reproduction on
subsurface rootstocks and its widespread
distribution in the western United States
(Foote et al. 1993), where alternate hosts
other than S. californica must occur, we be-
lieve that P. anthracina probably is nearly
monophagous on Solidago spp. of similar
growth habit. This assessment of its host
affinities discounts the published ‘‘unpub-
lished record”’ for an unidentified species of
Erigeron in Wasbauer (1972) cited by Foote
et al. (1993). Erigeron belongs to a differ-
ent subtribe, Asterinae, of the tribe Solida-
gininae, than Solidago (Bremer 1994).
Foote and Blanc (1963) reported a sweep
complex, 1—spiracle, 2—verruciform sensillum; (G) caudal segment, 1—posterior spiracular plates, 2—com-
pound sensillum, stelex sensillum, 3—compound sensillum, verruciform sensillum; (H) posterior spiracular plate,
1—rima, 2—interspiracular process.
186 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Bigs 3:
anterior view, |
dorsal sensory organ, 2—terminal sensory organ, 3—pit sensory organ, 4—lateral sensory organ, 5—supralateral
sensory organ; (D) caudal segment, 1—stelex sensillum, 2—rima, 3—interspiracular process, 4—compound
Second instar of Procecidochares anthracina. (A) habitus, anterior to left; (B) gnathocephalon,
anterior sensory lobe, 2—stomal sense organ, 3—mouth hooks; (C) anterior sensory lobe, 1—
sensillum, stelex sensillum, 5—compound sensillum, verruciform sensillum.
record from a Chrysothamnus sp., but that
plant is a doubtful host of P. anthracina.
Sweep records of adults all too often are
misleading and suspect indicators of the
host-plant affinities of nonfrugivorous te-
phritids. All four species of Procecidochar-
es from southern California previously
studied in similar detail as P. anthracina
(Silverman and Goeden 1980, Green et al.
1993, Goeden et al. 1994a, Goeden and
Teerink 1997) are strictly monophagous or
nearly monophagous on congeneric hosts.
BIOLOGY
Egg.—Galls of P. anthracina were found
on shoots arising from rhizomes of Soli-
dago californica under crowns of decidu-
ous oaks within a 3- to 10-cm deep surface
layer of humus, 1.e., decayed leaves, twigs,
branches, and acorns; therefore, females
must penetrate this loosely packed organic
layer to reach the buried axillary buds with-
in which the eggs are deposited (Fig. 6A,
B). This behavior would require that fe-
males climb down the buried parts of ver-
tical aerial shoots, or buried tree trunks,
near to which galls most commonly were
found. Females apparently may also follow
farther along the horizontal rhizomes at the
humus/mineral-soil interface and climb one
or more vertical subsurface shoots to ovi-
posit in the axillary buds. Alternatively, the
females may sense and reach these buried
shoots by penetrating the humus from di-
VOLUME 99, NUMBER 1
Fig. 4.
view, |—dorsal sensory organ, 2—anterior sensory lobe, 3—stomal sense organ; (C) anterior sensory lobe, 1—
dorsal sensory organ, 2—terminal sensory organ, 3—pit sensory organ, 4—lateral sensory organ, 5—supralateral
2—1rima, 3—interspiracular process.
sensory organ; (D) caudal segment, 1—stelex sensillum,
rectly above (like a pig rooting for truffles),
but this preovipositional searching behavior
seems less certain than using shoots and
rhizomes as pathways. Ovipositing adults
never were observed in nature, only in in-
sectary cages (Fig. 6A).
The eggs were inserted pedicel upward,
side-by-side, and lengthwise in bands with-
in a prominent axillary bud, but not pene-
trating any tissue. Instead, the eggs were
laid parallel to the long axis of the bud and
stem, between an outer bract and bud, be-
tween the bud and stem, or among the leaf
primordia (Fig. 6B). As many as 16 eggs
were found in a single bud in nature;
whereas, as many as 18 eggs were laid by
one female in | day in an axillary bud in
insectary cagings (Fig. 6A, B).
First instar of Procecidochares anthracina. (A) habitus, anterior to right; (B) gnathocephalon, anterior
Larva.—Eggs hatch about a week after
their deposition (Silverman and Goeden
1980, Green et al. 1993), but then the first
instars persist, as these and later instars de-
velop slowly through the fall, winter, and
spring. The first instars initially develop
gregariously by feeding within axils and
buds in shallow, linear, surface scars (Fig.
6C). Some first instars move to adjacent ax-
illary buds on the same shoot; others re-
main behind and feed gregariously within
the original bud or divide into subgroups
that subsequently each form separate galls
that may arise from one axillary bud. Gall
formation appears to result from larval ac-
tivities, not maternal stimuli, as bud and
stem swelling begins only after the larvae
begin their surface feeding. Multiple gall
188 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1SKV K226 6812
160, au
Fig. 5. Puparium of Procecidochares anthracina.
(A) habitus, anterior to left; (B) anterior end, 1—in-
vagination scar, 2—prothoracic rugose pads, 3—pro-
thoracic verruciform sensilla, 4—anterior spiracles;
(C) caudal segment, 1—posterior spiracular plates, 2—
compound sensillum.
formation from single aerial axillary buds
also was reported for P. kristinae and P.
lisae (Goeden and Teerink 1997).
Examination of 140 shoots on rhizome
fragments excavated at study site 1 on
8.x1i.1994, yielded seven shoots (5%) an
average of 10.2 + 0.6 (range, 1.9—23) mm
in length that together bore 14 axillary buds
and 16 + 2.5 (range 7—25) early first instars
per shoot. These infested axillary buds av-
eraged 0.56 + 0.06 (range, 0.37—1.12) mm
in length by 0.42 + 0.04 (range, 0.18—0.84)
mm in width and each contained an average
of 6 + 1.0 (range, 1—16) early first instars
(Fig. 6C). Infested buds were located an av-
erage of 3.9 + 0.6 (range, 1.8—8.0) mm be-
low the shoot apices (Fig. 6D).
Five infested shoots on five separate rhi-
zome fragments excavated at the same site
on 2.11.1995 each bore an average of 14 +
2 (range, 9-20) late first instars, or an av-
erage of 6 + 1.4 (range, 3-10) larvae per
axillary bud. The swollen axillary buds (in-
cipient galls) measured 2.56 + 0.67 (range,
0.74—4.56) mm long by 1.63 = 0.35 (range,
0.74—2.56) mm wide and contained sub-
spheroidal, open central cavities 0.2—0.4
mm in diameter (Fig. 6E). These incipient
galls were borne on shoots that averaged
9.4 + 1.4 (range, 6.6—13.7) mm in length
(Fig. 6D).
The only galls found containing second
instars were five in a sample collected at
site 1 on 28.i111.1994. The subspheroidal
galls contained an average of 4 + | (range,
2-8) second instars and averaged 3.9 + 1.4
(range, 1.5—9.1) mm in length by 5.9 + 2.0
(range, 1.9-10) mm wide. Each gall con-
tained a central cavity 1.1—4.0 mm in di-
ameter and had walls 1.6 + 0.2 (range, 1.3-
2.6) mm thick (Fig. 6F).
Galls containing third instars, or some-
times third instars and puparia (Fig. 6H),
were sampled more frequently than those
containing the first two instars at both study
sites. Twenty clusters of full-size galls as
shown in Fig. 6G contained an average of
6 + | (range, 1-14) galls and an average
total of 12 + 2 (range, 2—43) third instars
per shoot. These galls were subspheroidal,
ovoidal, or obclavoidal in shape and pale
violet basally, whitish medially, and pale
green or white apically (Fig. 6G). Forty-
a
VOLUME 99, NUMBER | 189
Fig. 6. Life stages of Procecidochares anthracina on Solidago californica. (A) ovipositing female, (B)
axillary bud on shoot with bracts removed to show newly deposited eggs (arrows), (C) first instar larvae (arrows)
feeding gregariously on shoot stem (line = 0.1 mm), (D) galled axillary bud on stalk of shoot dug from humus,
(E) sagittal section of gall containing second instars (arrows), (F) sagittal section of gall containing early third
instars, (G) cluster of full-size galls, (H) sagittal section through full-size gall containing third instars and puparia,
and (I) full-size gall showing exit channel and empty puparia. Lines = | mm, except as noted.
190 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
four (54%) of 81 galls with third instars
were borne on pedicels or stalks 7.7 + 1.0
(range, 2—34) mm in length; the remainder
(46%) were sessile on shoots. The 81 galls
averaged 13.3 + 0.3 (range, 8—20) mm in
length and 8.0 + 0.2 (range, 3.1-12) mm
in width. Each gall contained an average of
3 + 0.2 (range, 1—10) third instars within a
subspheroidal, ovoidal, or obovoidal, open
central cavity, which averaged 6.06 + 0.20
(range, 1.84—10.64) mm in length and 3.81
+ 0.12 (range, 1.68—6.72) mm in width.
Eighteen (22%) of these galls also bore an
average of 2 + 0.4 (range, 1—7) puparia in-
dicating differential rates of development
within galls. Another 145 galls were ex-
amined that only contained an average of 5
+ 0.4 (range, 1-21) empty (Fig. 6D or still
intact puparia.
Adult.—Eclosing adults emerge through
a common, open exit channel at the apex of
each gall, past and through an encircling
whorl of bracts, and continue upward
through the humus to reach the surface.
Adults were relatively long lived among
gallicolous congeners studied in southern
California to date (Silverman and Goeden
1980, Green et al. 1993, Goeden and Teer-
ink 1997). Twenty seven males lived an av-
erage of 47 + 3 (range, 14—70) days in in-
sectary cagings; 19 females averaged 38 +
3 (range, 14—62) days. Females (Fig. 7A)
are sexually immature at emergence (n =
3), but within a week produce a full com-
plement of mature ova (n = 3), apparently
mainly from tiny fat bodies (globules) ob-
served in the haemolymph of newly
emerged flies, which disappear during ovi-
genesis. In laboratory arenas, males 1-18
days old and females 1—22 days old mated
(Fig. 7B, C), but this behavior, like ovipo-
sition, never was observed in nature nor
were free-living adults ever collected.
The adults exhibited reproductive behav-
ior typical of other circumnatal, gallicolous
species of Procecidochares studied in
southern California, i.e., a lack of courtship
behavior, the exhibition of enantion type
wing movements by both sexes, and male
iG
Fig. 7. Procecidochares anthracina. (A) dorsal
view of female, (B) mating pair, lateral view, (C) Mat-
ing pair, ventral view. Lines = 1 mm.
stalking of females prior to mating (Green
et al. 1993, Headrick and Goeden 1994);
however, they also displayed several unique
behaviors in oviposition cages and petri
dish arenas. For example, females displayed
rapid enantion in brief spurts, two or three
times per second after they alternately or
synchronously supinated their wings
through arcs of 20-30° as they briefly
VOLUME 99, NUMBER 1
paused while walking along caged shoots
between ovipositions, or while resting after
ovipositing. So rapid was this enantion that
the wings became blurred. Four females
climbed atop prominent buds on excised
shoots during oviposition in cages, re-
curved their abdomens, and inserted their
aculei between bracts for an average of 6
+ 2 (range, 3—12) minutes, taking about 2
minutes to lay a single egg (Fig. 6A).
Both sexes displayed wing supination
blending into rapid wing enantion while
tracking each other in arenas prior to cop-
ulation, or rapid enantion during copulation
if agitated, especially the females, just prior
to and following postcopulatory separation.
Twenty seven copulations averaged 63 + 6
(range 25—148) minutes in duration and oc-
curred throughout the day, but mostly at
dusk. Males dismounted females either by
walking forward over the head of the fe-
males or by stepping off to one side of her
as she turned 180°, and they continued
walking in opposite directions, as the
male’s genitalia was pulled free; the entire
process lasted only a few seconds to as long
as 20 seconds (n = 4). The only other
unique behavior observed was between two
different pairs of males which stood facing
each other with their foretarsi overlapped
and touched mouthparts or one male
touched his labellum to the antennae and
frons of the other male. Nonreceptive fe-
males resisted the attentions of males by
head-butting; by turning swiftly towards
males attempting to mount them from the
rear, as is usual; and by not parting their
wings and to prevent newly mounted males
from grasping the females’ anterolateral ab-
dominal terga with their foretarsi.
Seasonal history.—Procecidochares an-
thracina is a univoltine, circumnatal species
(Headrick and Goeden 1994) on S. califor-
nica in southern California. The overwin-
tering stages are first- and second-instar lar-
vae which continue their slow, protracted
development beneath insulating snow and
humus as the winter progresses. When
spring arrives and their host plant resumes
19]
growth during April—May, the galls and lar-
vae within resume their growth and devel-
opment, the larvae pupariate, and the adults
emerge in late summer (July—September),
to mature, mate, and oviposit as described
above.
Natural enemies.—Two species of chal-
cidoid Hymenoptera have been reared by us
as parasitoids of P. anthracina. Eurytoma
obtusiventris Gahan (Eurytomidae) is a sol-
itary, larval-pupal endoparasitoid recovered
from puparia that also has been reported
from the tephritids, Eurosta_ solidaginis
(Fitch) and Neotephritis (as Tephritis) fin-
alis (Loew) (Bugbee 1967, Foote et al.
1993). Pronotalia carlinarum Gradwell
(Eulophidae) is a gregarious puparial en-
doparastoid, the only species in this genus
known from North America, and a known
gregarious endoparasitoid of puparia, main-
ly Tephritidae, but also Agromyzidae and
Chloropidae (LaSalle 1994).
ACKNOWLEDGMENTS
We sincerely thank A. C. Sanders, Cu-
rator of the Herbarium, Department of Bot-
any and Plant Sciences, University of Cal-
ifornia, Riverside, for identification of
plants. The parasitoids were identified by
Michael Gates, Department of Entomology,
University of California, Riverside. We also
are grateful to K L. Blanc, D. H. Headrick,
A. L. Norrbom, and G. J. Steck for their
helpful comments on earlier drafts of this
paper.
LITERATURE CITED
Bezzi, M. 1910. Restaurazione del genere Carpomyia
(Rond.) A. Costa. Bollettino del Laboratorio di
Zoologia generale e agraria della R. scuola supe-
riore d’agricoltura in Portici 5: 3-33.
Bezzi, M. and J. S. Tavares. 1916. Alguns muscideos
cecidogenicos do Brasil. Broteria (Serie Zoologi-
ca) 14: 155-170.
Bremer, K. 1994. Asteraceae Cladistics & Classifica-
tion. Timber Press, Inc. Portland, Oregon.
Bugbee, R. E. 1967. Revision of chalcid wasps of
genus Eurytoma in America North of Mexico.
Proceedings of the United States. National Mu-
seum 118: 433-552.
Doane, R. W. 1899. Notes on Trypetidae with descrip-
192 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tions of new species. Journal of the New York
Entomological Society 7: 177-193, pl.
Foote, R. H. and FE L. Blanc. 1963. The fruit flies or
Tephritidae of California. Bulletin of the Califor-
nia Insect Survey 7, 117 p.
Foote, R. H., EF L. Blanc, and A. L. Norrbom. 1993.
Handbook of the Fruit Flies (Diptera: Tephritidae)
of America North of Mexico. Cornell University
Press, Ithaca, New York.
Goeden, R. D. and D. H. Headrick. 1990. Notes on
the biology and immature stages of Stenopa affinis
Quisenberry (Diptera: Tephritidae). Proceedings
of the Entomological Society of Washington 92:
641-648.
. 1991a. Notes on the biology, hosts, and im-
mature stages of Tomoplagia cressoni Aczél in
southern California. Proceedings of the Entomo-
logical Society of Washington 93: 549-558.
1991b. Life history and descriptions of im-
mature stages of Tephritis baccharis (Coquillett)
on Baccharis salicifolia (Ruiz & Pavon) Persoon
in southern California (Diptera: Tephritidae). Pan-
Pacific Entomologist 67: 86—98.
1992. Life history and descriptions of im-
mature stages of Neaspilota viridescens Quisen-
berry (Diptera: tephritidae) on native Asteraceae
in southern California. Proceedings of the Ento-
mological Society of Washington 94: 59-77.
Goeden, R. D., D. H. Headrick, and J. A. Teerink.
1993. Life history and descriptions of immature
stages of Tephritis arizonaensis Quisenberry
(Diptera: Tephritidae) on Baccharis sarothroides
Gray in southern California. Proceedings of the
Entomological Society of Washington 95: 210—
220
. 1994a. Life History and descriptions of im-
mature stages of Procecidochares flavipes Aldrich
(Diptera: Tephritidae) on Brickellia spp. in south-
ern California. Proceedings of the Entomological
Society of Washington 96: 288-300.
1994b. Life history and description of im-
mature stages of Paroxyna genalis (Thomson)
(Diptera: Tephritidae) on native Asteraceae in
southern California. Proceedings of the Entomo-
logical Society of Washington 96: 612-629.
1995a. Life history and description of im-
mature stages of Urophora timberlakei Blanc and
Foote (Diptera: Tephritidae) on native Asteraceae
in southern California. Proceedings of the Ento-
mological Society of Washington 97: 780-791.
1995b. Life history and descriptions of im-
mature stages of Valentibulla californica (Coquil-
lett) (Diptera: Tephritidae) on Chrysothamnus
nauseosus (Pallas) Britton in southern California.
Proceedings of the Entomological Society of
Washington 97: 548-560.
Goeden, R. D. and J. A. Teerink. 1996a. Life histories
and descriptions of adults and immature stages of
two cryptic species, Aciurina ferrugenia (Doane)
and A. michaeli new species (Diptera: Tephritidae)
on Chrysothamnus viscidiflorus (Hooker) Nuttall
in southern California. Proceedings of the Ento-
mological Society of Washington 98: 415—438.
. 1996b. Life history and descriptions of adults
and immature stages of Aciurina idahoensis
Steyskal (Diptera: Tephritidae) on Chrysothamnus
viscidiflorus (Hooker) Nuttall in southern Califor-
nia. Proceedings of the Entomological Society of
Washington 98: 681-694.
. 1996c. Life history and descriptions of adults
and immature stages of Aciurina semilucida
(Bates) (Diptera: Tephritidae) on Chrysothamnus
viscidiflorus (Hooker) Nuttall in southern Califor-
nia. Proceedings of the Entomological Society of
Washington 98: 752-766.
. 1997. Notes on life histories and descriptions
of adults and immature stages of Procecidochares
kristineae and P. lisae new spp. (Diptera: Tephrit-
idae) on Ambrosia spp. in southern California.
Proceedings of the Entomological Society of
Washington 99: 67-88.
Green, J. FE, D. H. Headrick, and R. D. Goeden. 1993.
Life history and description of immature stages of
Procecidochares stonei Blanc & Foote on Vigui-
era spp. in southern California (Diptera: Tephrit-
idae). Pan-Pacific Entomologist 69: 18-32.
Headrick, D. H. and R. D. Goeden. 1990. Description
of the immature stages of Paracantha gentilis
(Diptera: Tephritidae). Annals of the Entomolog-
ical Society of America 83: 220-229.
1991. Life history of Trupanea californica
Malloch (Diptera: Tephritidae) on Gnaphalium
spp. in southern California. Proceedings of the En-
tomological Society of Washington 93: 559-570.
. 1993. Life history and description of imma-
ture stages of Aciurina thoracica (Diptera: Te-
phritidae) on Baccharis sarothroides in southern
California. Annals of the Entomological Society
of America 86: 68-79.
Headrick, D. H., R. D. Goeden, and J. A. Teerink.
1995. Life history and description of immature
stages of Euaresta stigmatica Coquillett (Diptera:
Tephritidae) on Ambrosia spp. (Asteraceae) in
southern California. Annals of the Entomological
Society of America 88: 58-71.
. 1996. Life history and description of imma-
ture stages of Dioxyna picciola (Bigot) (Diptera:
Tephritidae) on Coreopsis spp. (Asteraceae) in
southern California. Proceedings of the Entomo-
logical Society of Washington 98: 332-349.
Knio, K. M., R. D. Goeden, and D. H. Headrick. 1996.
Descriptions of immature stages of Trupanea ni-
gricornis and T. bisetosa (Diptera: Tephritidae).
Annals of the Entomological Society of America
89: I-11.
LaSalle, J. 1994. North American genera of Tetrasti-
VOLUME 99, NUMBER 1 198
chinae (Hymenoptera: Eulophidae). Journal of ifornia (Diptera: Tephritidae). Pan-Pacific Ento-
Natural History 28: 109-236. mologist 56: 283-288.
Munz, P. A. 1974. A Flora of Southern California. | Wasbauer, M. W. 1972. An annotated host catalog of
University of California Press, Berkeley. the fruit flies of America north of Mexico (Dip-
Silverman, J. and R. D. Goeden. 1980. Life history tera: Tephritidae). California Department of Ag-
of a fruit fly, Procecidochares sp., on the ragweed riculture, Bureau of Entomology, Occasional Pa-
Ambrosia dumosa (Gray) Payne, in southern Cal- pers 19: 1-72.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 194—203
A REVISION OF THE NEOTROPICAL ANTS OF THE GENUS
CAMPONOTUS, SUBGENUS MYRMOSTENUS
(HYMENOPTERA: FORMICIDAE)
WILLIAM P. MACKAY
Laboratory for Environmental Biology, Centennial Museum, The University of Texas,
El] Paso, TX 79968, U.S.A.
Abstract.—The subgenus Myrmostenus consists of a group of 6 species of South Amer-
ican (Peru, Bolivia and Brasil) ants which are known only from the females. The species
include C. convexiclypeus MacKay (new species), C. leptocephalus Emery, C. longipilis
Emery, C. mirabilis Emery, C. postangulatus Emery (new status) and C. sphenocephalus
Emery. They are easily recognized as the head is greatly elongated and somewhat flat-
tened. Nothing is known of the biology of this interesting subgenus. A key is provided
for identification of the species.
Resumen.—El subgénero Myrmostenus consiste de un grupo de seis especies de hor-
migas de Pert: y Brasil conocidas solamente por las hembras. Las especies incluyen C.
convexiclypeus MacKay, C. longipilis Emery, C. leptocephalus Emery, C. mirabilis Em-
ery, C. postangulatus Emery y C. spenocephalus Emery. Se pueden reconocer porque la
cabeza es elongada y aplanada. La biologia de estas hormigas interesantes es desconocida.
Se incluye una clave para la determinacion de las especies.
Key Words:
Peru, Brasil
With the exception of the small Nearctic
and European faunas, the genus Campono-
tus currently consists of poorly defined
groups (“‘subgenera’’) in which species
identification is nearly impossible. The ge-
nus as a whole is often considered an enor-
mous group of rather bland ants, with few
good characters for separating species.
While this is probably true for some sub-
genera (1.e., Tanaemyrmex, Mymobrachys),
there are certainly many exceptions. One of
the most fascinating groups in the genus is
the subgenus Myrmostenus which, unfor-
tunately, is known only from the females.
Species in this subgenus are easily recog-
Neotropics, Peru, Brasil, Camponotus, Myrmostenus, Areas neotropicales,
nized by their distinctive, elongate heads
and, as this study reveals, have good char-
acters for the separation of species. Nothing
is known of the biology of this curious
South American subgenus.
This is the first in a series of contribu-
tions towards the understanding of the New
World species of this large, complex genus.
MATERIALS AND METHODS
Specimens were borrowed from several
institutions and curators as follows:
AMNH, American Museum of Natural
History, New York, Mark Smethurst.
CASC, California Academy of Sciences,
VOLUME 99, NUMBER 1
San Francisco, Darrell Ubick, Wojciech Pu-
lawski.
LACM, Los Angeles County Museum,
Los Angeles, Roy Snelling.
MCSN, Museo Civico di Storia Naturale,
Valter Raineri (Emery Collection), Genoa,
Italy.
MCZC, Museum of Comparative Zool-
ogy, Harvard University, Cambridge, Mas-
sachusetts, Stefan Cover.
USNM, National Museum of Natural
History, Smithsonian Institution, Washing-
ton, D.C., David Smith.
Specimens were measured using an oc-
ular micrometer in a dissecting microscope.
The following abbreviations are used (all
measurements in mm.):
HL, Head length, anterior of median lobe
of clypeus to mid point of occiput (ex-
cluding occipital corners).
HW, Head width, maximum excluding eyes
(immediately anterior to eyes).
EL, Eye length, maximum dimension.
SL, Scape length, excluding basal condyle.
Clypeal length, Distance from the tops of
the lobes of the posterior border of the
clypeus to the edge of the anterior medial
border (Fig. 8a).
Clypeal width, Distance between the ten-
torial pits (Fig. 6b).
Clypeal index, Width/length x 100.
CI, Cephalic Index, HW/HL xX 100.
SI, Scape index, SL/HL X_100 (mote: HL
used instead of HW).
SUBGENUS MYRMOSTENUS
Camponotus subgenus Myrmamblys (in
part): Forel, 1914:271, 272
Camponotus subgenus Myrmomalis (in
part): Forel, 1914:271
Camponotus subgenus Myrmostenus Em-
eny, 1920:250"260
Camponotus subgenus Pseudocolobopsis
(in part): Emery, 1923:62
Type species of the subgenus Myrmostenus:
Camponotus mirabilis Emery, 1903 (des-
ignated by Emery, 1920)
Diagnosis.—The females of this subge-
nus have greatly elongated heads with a CI
195
ranging from 48—101. The pronotum is also
long, together with the remainder of the
mesosoma. The basal face of the propo-
deum is much shorter than the dorsal face.
Most surfaces are smooth and _ shining.
Erect hairs are usually sparse and decum-
bent pubescence is usually absent. The head
has a few long, coarse hairs near the top of
head and usually along the frontal carinae.
The cheeks extend past the base of the man-
dibles, usually forming distinct lobes. The
workers and males are unknown.
Description.—Female measurements
(mm): HL 2.36—4.70, HW 1.44—2.52, SL
208—3.76, EL 0.56—0.98. Indices: SI 48-
101, Cl 40-67.
Mandible with 6 poorly defined teeth, ex-
cept for apical and subapical; clypeus large
and wide, often with well defined median
carina; clypeal border slightly concave in
all species except C. mirabilis; eyes large
and extending past lateral edges of head;
ocelli well developed; scape extending past
posterior border of head (except in C. mi-
rabilis); vertex concave or convex; maxil-
lary palps with 6 segments; labial palps
with 4 segments; mesosoma somewhat
elongated and flattened; pronotum especial-
ly elongate; propodeum with long basal
face and very short posterior face; petiole
small, petiolar node relatively sharp.
Hairs sparse and coarse (except for C.
leptocephalus, which has abundant erect
hairs); decumbent pubescence essentially
absent (except C. leptocephalus, which has
sparse pubescence).
Sculpture fine, most surfaces smooth and
polished, except head, which is punctate in
most species.
Color medium brown.
Worker and Male: Unknown
Distribution.—Peru, Bolivia and Brasil.
Discussion.—These ants are consider-
ably modified, possibly for living under
bark or in tunnels in wood. It is difficult to
determine the affinities with other subgen-
era in Camponotus, but the lobes associated
with the cheeks, the enlarged clypeus and
polished integument suggests that they may
196 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
be related to Pseudocolobopsis. They are,
however, much larger than any known
Pseudocolobopsis females.
Biology.—Unknown.
SPECIES COMPLEXES
This subgenus contains three species
complexes, the /eptocephalus complex, the
longipilis complex and the mirabilis com-
plex. The leptocephalus complex consists
of a single species, C. leptocephalus. This
complex can be characterized as having a
relatively larger CI (61), and having most
surfaces covered with short, bristly hairs.
The sculpture is rougher than in other spe-
cies in the subgenus, especially obvious on
the mesosoma. The lJongipilis species com-
plex consists of four species: C. convexicly-
peus, C. longipilis, C. postangulatus and C.
sphenocephalus. These species can be char-
acterized by having intermediate values of
CI (51-67), lacking bristly hairs and having
smooth and polished sculpture on the me-
sosoma and gaster, with the head sculptured
to varying degrees with punctures. Finally,
the mirabilis complex consists of a single
species, C. mirabilis. This complex is char-
acterized by having a very long head (CI
40—44), which is noticeably widened ante-
riorly, lacking short, bristly hairs and in be-
ing smooth and shining throughout, includ-
ing the head.
I am not interested in defending the con-
cept of subgenera as valid taxonomic
groupings, but only as a groupings of con-
venience. In this instance, we can group
three apparently related species complexes
conveniently into a larger group, the sub-
genus Myrmostenus.
KEY TO THE FEMALES OF THE ANTS OF THE
SUBGENUS MyYRMOSTENUS
1. Ventral surface of head with more than 5
erect hairs (Figs. 1, 2); occipital corners of
head rounded (Figs. 4, 7, 8)
— Ventral surface of head without erect hairs,
or rarely with 1 or 2 present; occipital cor-
ners of head angulate (Figs. 5,6) ........ 5)
2(1). Surface of clypeus with more than 20 erect
nen (ENS. (3) 5 5 6 o> oo ec leptocephalus Emery
— Surface of clypeus usually without any erect
hairs (Fig. 7), although there may be occa-
sional erect hairs along the borders (Fig. 5)
3(2). Head strongly narrowed posteriorly, the
minimum posterior width less than the dis-
tanceybetweenveyes) (Eicss7) ee eae
se tee Epes ae ele sphenocephalus Emery
— Head not strongly narrowed posteriorly
(Figs. 4, 8), or minimum posterior width
much greater than distance between eyes,
nearly as wide as width of head anterior to
eyes (Fig. 6)
4(3). Head relatively wide, CI > 60 (Fig. 4);
scape long, SI > 85; eye large, maximum
diameter > 0.85mm ... . convexiclypeus, n.sp.
— Head narrowed, CI < 60 (Fig. 6); scape
shorter, SI < 85; eye diameter smaller, max-
imum diameter < 0.85mm_ . . longipilis Emery
5(1). Head greatly elongate (CI 40—44) with
strongly formed occipital angles (Fig. 5); rel-
atively commonly collected .. mirabilis Emery
— Head not greatly elongate (CI 55), occipital
angles poorly formed (Fig. 6); rarely col-
lected hee eae - postangulatus Emery
CLAVE PARA LA DETERMINACION DE LAS
HEMBRAS DEL SUBGENERO MYRMOSTENUS
ile Superficie ventral de la cabeza con 5 0 mas
pelos rectos (Figs. 1, 2); esquinas occipitales
de la cabeza redondas (Figs. 4, 7, 8) ..... 2)
— Superficie ventral de la cabeza sin pelos rec-
tos; esquinas occipitales anguladas (Figs. 5,
GYR HU TE Te We ene 5
2(1). Superficie del clipeo con 20 o mas pelos
Mes ENS) oso55scc leptocephalus Emery
— Superficie del clipeo usualmente sin pelos
rectos (Fig. 7), aunque podria tener pelos
rectosyalemareenu(Eiese) eee eine eee 3
3(2). Cabeza muy angosta atras, la anchura pos-
terior menos que la distancia entre los ojos
(Eis ye eee, ee ee sphenocephalus Emery
— Cabeza no muy angosta atras (Figs. 4, 8), o
la anchura posterior casi tan ancha como la
anchura inmediamente detras de los ojos
(EVD31@) eg teg oth eee rs eee longipilis Emery
4(3). Cabeza no muy angosta (indice cefalico >
60, Fig. 4); escapo largo (indice del escapo
> 85); diametro maximo del ojo > 0.85mm
Aa opted oa iacg Maecsteay ena hee convexiceps, N.sp.
— Cabeza angosta (indice cefalico < 60, Fig.
6); escapo mas corto (indice del escapo <
85); diametro maximo del ojo < 0.85mm
Si tones AE longipilis Emery
5(1). Cabeza muy elongada (CI 40—44), con 4n-
gulos bien formados en las esquinas occip-
itales (Fig. 5); relativemente comun ...
Bi) TEER Ode LP nee ee ee aes mirabilis Emery
VOLUME 99, NUMBER 1
— Cabeza no demasiada elongada (CI 55), con
Angulos poco diferenciados en las esquinas
occipitales (Fig. 6); no comunes......
if Nina ck UNECE ero postangulatus Emery
DESCRIPTIONS OF SPECIES
Camponotus (Myrmostenus)
convexiclypeus MacKay, new species
(Figs. 3, 4; Map 2)
Diagnosis.—This is a large species with
a head much wider than in the other species
in the subgenus (2.52 mm vs. 1.44—2.00
mm in all other species in the subgenus).
The clypeus is strongly convex as seen
from above (Fig. 3). It has the usual coarse
hairs on the vertex and also on the under-
side of the head.
Description.—Female measurements
(nm): 3H 3.76; HW 2:52; SE. 3.76, EL
0.98, clypeus length 1.52, clypeus width
1-22. Indices: SI 100; Cl 67,,clypeal imdex
80.
Mandible with 6 teeth and an angle at
basal margin; clypeus longer than wide,
strongly convex (Fig. 3); anterior clypeal
border concave with two lateral lobes;
scape extending about 2 length past pos-
terior border of head; head flattened in pro-
file; vertex convex; maxillary palps short,
about 4 length of distance to foramen mag-
num; labial palps also short, about % length
of maxillary palps; mesosoma lengthened
as in other species; propodeum with poorly
defined angle between two faces, posterior
face appearing much shorter than basal
face; petiole thin, with convex anterior face
and straight posterior face (Fig. 2).
Hairs sparse, but coarse, some bent at an-
gles and long (0.70—0.90 mm), area from
vertex to upper edge of frontal carinae with
about 20 coarse hairs, clypeus without hairs
except for fringe at anterior border, about
20 hairs on underside of head, scapes with
scattered erect hairs, up to 0.25 mm in
length, dorsum of mesosoma with about 20
hairs, propodeum at point where faces meet
with 6 hairs on left side, three on right side,
node of petiole with three hairs on left, one
on right, gaster with scattered hairs at the
197
edges of each tergum but also on the other
surfaces; decumbent pubescence almost ab-
sent.
Sculpture coarse on head, consisting of
dense punctures, similar to that of C. sphen-
ocephalus, remainder of body smooth and
shining.
Color medium brown, head, including
mandibles, scapes and funiculi, darker
brown.
Type series.—Holotype 2°, PERU, Mon-
son Valley, Tingo Maria, 11-x1i-1954; E. I.
Schlinger & E. S. Ross collectors.
Material examined.—Holotype @&@
(CASC).
Distribution—Known only from type lo-
cality.
Etymology.—Refers to the strongly con-
vex clypeus.
Discussion.—This species appears to be
related to C. sphenocephalus, as the vertex
is rounded and the head is densely punctate
as in C. sphenocephalus. The head is wider
as seen in full face view, thus making it
easily separated from C. sphenocephalus.
Biology.—Unknown.
Camponotus (Myrmostenus) leptocephalus
Emery
(Figs. 2, 3, 8; Map 1)
Camponotus (Pseudocolobopsis) leptoceph-
alus Emery, 1923:62, female, Fig. 3,
BRASIL, Espirito Santo, MCSN [seen].
Diagnosis.—This species has the least
elongate head among the species in this
subgenus (CI 61). It can be easily distin-
guished as it is much more hairy, with the
hairs being primarily short bristles. The
sculpture is relatively more rough than in
the other species in the complex.
Description.—Female measurements
(mm): Hi 2.36, HW 1.44, SL 2.38, EL
0.56, clypeal length 0.74, width 0.70. In-
dices: SI 101, CI 61, clypeal index 95.
Mandible with 6 teeth; clypeus slightly
longer than broad, with poorly defined me-
dian carina, anterior clypeal border slightly
concave; scape long, extending nearly half
198 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
|
es
Figs. 1-8. Females of Camponotus (Myrmostenus); scales are 0.2 mm. 1, C. (M.) longipilis, lectotype from
the side. 2, C. (M.) leptocephalus, \lectotype head, mesosoma, and petiole. 3, Clypeus of the 6 species, as seen
from the top. 4, Head of C. (M.) convexiclypeus, holotype. 5, C. (M.) mirabilis, head of lectotype. 6, C. (M.)
postangulatus, holotype head (b indicates clypeal width measurement). 7, C. (M.) spenocephalus, holotype head.
8, C. (M.) leptocephalus, lectotype head (a indicates clypeal length measurement).
Figs. 1-8. Hembras de Camponotus (Myrmostenus); las escalas son de 0.2 mm. 1, C. (M.) longipilis lectotipo
desde el lado. 2, C. (M.) leptocephalus cabeza, mesosoma y peciolo del lectotipo. 3, Clypeus de las 6 especies.
4, Cabeza de C. (M.) convexiclypeus holotipo. 5, C. (M.) mirabilis cabeza del lectotipo. 6, C. (M.) postangulatus
cabeza del holotipo. 7, C. (M.) spenocephalus cabeza del holotipo. 8, C. (M.) leptocephalus cabeza del lectotipo.
its length past posterior border of head; maxillary palps extending about %4 distance
head slightly flattened when seen in profile; to foramen magnum; labial palps about %
vertex concave when seen from the front length of maxillary palps; mesosoma
and above, convex as seen in full face view; lengthened as in other species; propodeum
VOLUME 99, NUMBER 1 199
) OUTH AMER)
a 50 TH AMERICA
eee Venezuela
SOUTH AMERICA
oO
_— Venezuela
|
@ /Jepto- ®@ /ongipilis
cephalus B convex!-
BH post - clypeus
angulatus
UTH AMERICA
a clo
e)
_— Venezuela
Oe
mirabilis
spheno-
cephalus
Maps 1—4. Distribution of Camponotus (Myrmostenus). 1, C. (M.) leptocephalus and C. (M.) postangulatus.
2, C. (M.) longipilis and C. (M.) convexiclypeus. 3, C. (M.) mirabilis. 4, C. (M.) spenocephalus.
200 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
with posterior face about % of basal face;
petiole with convex anterior face, nearly flat
posterior face.
Hairs (erect and suberect) abundant on
head and gaster, few scattered erect hairs on
mesosoma and propodeum; decumbent pu-
bescence scattered, but sparse on all body
surfaces, length up to 0.06 mm in length.
Lightly but densely and evenly punctate
on head, coriaceous on all surfaces of me-
sosoma and gaster.
Color concolorous medium brown, gaster
slightly darker.
Type series.—Lectotype ¢ [here desig-
nated], Espir. Santo, X.1920-II.1921; Arp
dedic; Camp. leptocephalus n. (MCSN)
[seen]; second female not seen.
Material examined.—Lectotype °.
Distribution.—Known only from type lo-
cality.
Discussion.—This species forms one of
the extremes in the range of head shape,
sculpture and hairiness, and thus belongs to
a separate species complex. The head is not
as elongate as most of the other species (CI
61) or flattened as the other species. It ap-
pears to be distantly related to C. spheno-
cephalus, and can be easily separated as the
clypeus is covered with erect hairs (none in
C. sphenocephalus). This species is defi-
nitely not a member of the subgenus Pseu-
docolobopsis, as the maxillary palps are
long, which is not found in species in Pseu-
docolobopsis.
Biology.—Unknown.
Camponotus (Myrmostenus) longipilis
Emery
(Figs. 1, 3; Map 2)
Camponotus longipilis Emery, 1911:224,
female, PERU, WHudnuco: Pachitea
(MCSN) [seen].
Camponotus (Myrmamblys) longipilis: For-
el, 1914:271.
Camponotus (Myrmostenus) longipilis: Em-
ery, 1920:260.
Diagnosis.—This species has an elongate
head (CI 54—56) which is basically rectan-
gularly shaped. It is identical to that of C.
postangulatus (Fig. 6), except it lacks the
occipital angles. The ventral surface of the
head is covered with dense, long, erect,
twisted hairs. The occipital corners are
rounded and not angulate.
Description.—Female measurements
(mm) : HL 3.44—3.58, HW 1.88-1.94, SL
2.68—2.88, EL 0.73-0.79, clypeal length
1.23-1.32, clypeal width 0.91—1.02. Indi-
ces: SI 78-84, CI 54—56, clypeal index 74-—
TAE
Mandible with 6 teeth; clypeus longer
than broad, widened anteriorly, surface
strongly convex with well defined carina;
clypeal border strongly concave; scape ex-
tending past posterior border of head; ver-
tex weakly convex; maxillary palps short
and delicate, extending about half distance
to foramen magnum; labial palps also slen-
der and about % length of maxillary palps;
descending face of propodeum somewhat
more than % length of basal face; petiole
with strongly convex anterior face, straight
posterior face.
Hairs erect, sparse, long and coarse on
head (including underside of head), scat-
tered on mesosoma and scattered on gaster;
decumbent pubescence very fine and sparse
on mesosoma and gaster.
Head weakly, but densely punctate, me-
sosoma and gaster coriaceous, but strongly
shining.
Color medium brown, head, mandibles
and scape darker brown.
Type series.—Lectotype 2 (here desig-
nated), Pachita, Peru, Stdg; longipilis Em-
ery (MCSN) [seen]; second female men-
tioned by Emery (1911) not seen.
Material examined.—23 @, including
BOLIVIA: HuachiBeni, ix-1921, W. Mann
(1 ¢ USNM). PERU: Pachita, Marcapata,
Staudinger (lectotype 2 MCSN, CoType #
21592, MCZC, part of type series of C. mi-
rabilis), Madre de Dios, Rio Tambopata Re-
serve, 30 air km SW Pto. Maldonado,
290m, 6/25-xi-1979, J. B. Heppner (18 @
USNM), Cuzco, Pilcopata, 11/14-x1i-1979,
600m, J. B. Heppner (2 2 USNM).
es
VOLUME 99, NUMBER |
Distribution.—Peru and adjacent Bolivia.
Discussion.—This species would not be
confused with any other in the Jongipilis
complex, except C. postangulatus. It differs
in that the occipital corners are not angulate
and the ventral surface of the head is cov-
ered with long, coarse, erect hairs. The
sculpture is similar to that of C. leptoceph-
alus, but it lacks the abundant hairs on the
dorsum of the head, having only a few scat-
tered, coarse, erect hairs. The clypeal carina
is also much more differentiated than it is
in C. leptocephalus.
Biology.—Unknown.
Camponotus (Myrmostenus) mirabilis
Emery
(Figs. 3, 5; Map 3)
Camponotus mirabilis Emery, 1903:80,
Fie. 15, female, PERU, Vilcanota
(MCSN) [seen].
Camponotus (Myrmomalis) mirabilis: For-
el; 1914:271-.
Camponotus (Myrmostenus) mirabilis: Em-
ery, 1920:260.
Diagnosis.—This is the most common
species in the subgenus, and is also easily
differentiated from all others. The head is
greatly elongated and noticeably widened
anteriorly. The occipital corners are strong-
ly angulate as seen in full face view. The
clypeus is nearly flat, with little evidence of
a raised region in the area of the clypeal
carina. All of the surfaces, including the
dorsum of the head, are strongly polished.
Description.—Female measurements
(mm): HL 4.18—4.70, HW 1.82-1.88, SL
2.08-2.24, EL 0.66—0.70, clypeal length
1.19-1.28, clypeal width 0.98—1.08. Indi-
ces: SI 48-50, CI 40—44, clypeal index 82-—
84.
Mandible with apical and subapical teeth
large, well defined, at least 4 additional
teeth defined to various degrees; clypeus
weakly convex with little evidence of clyp-
eal carina as slightly raised strip; clypeal
border convex and rounded; scape short and
not reaching posterior border of head; head
201
more than twice as long as wide, noticeably
widened near mandibles; vertex strongly
concave, with occipital corners strongly an-
gulate; maxillary palps very short, barely
extending past buccal region; labial palps
nearly as long as maxillary palps; propo-
deum with descending face about half
length of basal face; petiole with strongly
convex anterior face, nearly flat posterior
face, thicker and less in height than in the
other species.
Hairs erect, long, coarse and sparse on
dorsum of head, pronotum, scutum and scu-
tellum, propodeum, node of petiole and
gaster; decumbent pubescence very weak
and sparse on most surfaces.
Sculpture weak, shiny and polished on
most surfaces.
Color medium brown, head, mandibles
and scape somewhat darker, gaster with yel-
low blotches on both sides of anterior sec-
tion of terga.
Type series.—Lectotype 2 [here desig-
nated], Vilcanota, Peru, Stdg; Camponotus
mirabilis n. sp. (MCSN) [seen]; Marcapata,
Peru; 2 paralectotypes [here designated],
#21592 (MCZC 2 &, third female with
same numbers and labels is C. longipilis).
Material examined.—105 2, including
BOLIVIA: Rio Benrf, Rurrenbaque, W.
Mann, Nov, 1921, Mulford Exped. (2 &
USNM). PERU: Maracapata (3 type &
MCSN, MCZC), Vilcanota and Marcapata;
Loromayu, 400-500 m, ix-62, L. Pena (3
2 LACM, | 2 MCZC), Cuzco Dept., Quin-
cemil, 750m, 14/31-viii-62, Pena (2 &
MCZC), Pilcopata, 8/10-xii-1979, J. B. He-
ppner, (10 2 USNM), Madre de Dios, Cuz-
co Amazonico, 15K NE of Puerto Maldo-
nado, S..Cover & J. Yobin, 23-vi-91 (ik 2
MCZC), Madre de Dios, Avispas, 400m,
1/15-x-62, Pena © 92% MCZC), Avispas,
10/30-ix-62, Pena (2 2 MCZC); in car Pu-
callpa X Aguaytia, 18-vii-1968, C. & B.
O’Brian (1 2 CASC), Rio Tambopata Re-
serve, 30 air KSW Puerto Maldonado,
12°50’S_ 69°20'W, 1/26-xi-1982, E. Ross
(23 2 CASC), same locality, 1/4-v-1984,
W. Pulawski (4 2 CASC), same locality,
202
290m, 2/30-xi-1979, J. B. Heppner, (48 &
USNM).
Distribution.—Southern Peru, Bolivia,
and adjacent western Brasil.
Discussion.—This species is at the other
extreme from C. leptocephalus in terms of
its morphology. The head is startling in
view, greatly elongated and slender. The
ventral surface of the head lacks erect hairs.
The entire surface is shiny. The occipital
angles are well differentiated from the re-
mainder of the head. This is one of the most
fascinating ants from the Neotropical re-
gion, a true jewel in the genus Camponotus.
Biology.—Unknown. This species is rel-
atively common. A rather minor effort in
Peru would result in the collection of work-
ers and males and reveal what would be
expected to be an interesting story about
this unusual species.
Camponotus (Myrmostenus) postangulatus
Emery, new status
(Fig. 6; Map 1)
Camponotus longipilis var. postangulata
Emery, 1911:225, female, PERU, Hua-
nuco: Pachitea (MCSN) [seen].
Camponotus (Myrmostenus) longipilis vat.
postangulatus: Emery, 1925:161.
Diagnosis.—This species is very similar
to C. longipilis, but can be separated easily
as the occipital angles are well differenti-
ated, and there are no erect hairs on the
ventral surface of the head. The surface of
the clypeus is strongly convex with a well
differentiated clypeal carina.
Description.—Female measurements
Guim): HIS 62, EW 2.00; SE 2:84, EL
0.76, clypeal length 1.30, clypeal width
0.96. Indices: SI 78, CI 55, clypeal index
74.
Mandible with 6 poorly defined teeth;
clypeus strongly convex with well devel-
oped carina; clypeal border weakly con-
cave; scape extending past posterior border
of head; head somewhat flattened when
viewed in profile, shape identical to that of
C. longipilis, except for presence of occip-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ital angles; vertex concave, with well de-
veloped occipital angles; maxillary and la-
bial palps as in C. longipilis; mesosoma and
petiole as in C. longipilis.
Hairs; decumbent pubescence and sculp-
ture as in C. longipilis.
Type series.—Holotype °, Pachitea,
Peru, Stdg; longipilis var. postangulata Em-
ery (MCSN) [seen].
Material examined.—Holotype °.
Distribution.—Central Peru.
Discussion.—This species is closely re-
lated to C. longipilis, but is distinct enough
to be regarded as a valid species. In addi-
tion, the type locality for both species is the
same. The characters in the key and in the
diagnoses of the two species should be suf-
ficient to separate the two species.
Biology.—Unknown.
Camponotus (Myrmostenus)
sphenocephalus Emery
(Fig. 4; Map 4)
Camponotus sphenocephalus Emery, 1911:
225, female, PERU, Hudanuco: Pachitea
(MCSN) [seen].
Camponotus (Myrmamblys) sphenocepha-
lus: Forel, 1914:272.
Camponotus (Myrmostenus) sphenocepha-
lus: Emery, 1920:260.
Diagnosis.—This species has a strongly
and densely punctate head, a strongly con-
vex clypeus with well developed median
carina, with numerous erect, coarse hairs on
the underside of the head. The head is
strongly narrowed posteriorly, with a
rounded vertex. It is not closely related to
any of the other known species, but is most
similar to C. longipilis and C. postangula-
tus.
Description.—Female measurements
(mm): HL 3.51-3.66, HW 1.78—1.96, SL
2.86—-3.06, EL 0.78—0.81, clypeal length
1.34—1.36, clypeal width 0.93—0.95. Indi-
ces: SI 80-84, CI 51-54, clypeal index 68—
TO:
Mandible with 6 teeth; clypeus strongly
convex with well defined carina; anterior
VOLUME 99, NUMBER 1
clypeal border concave; scape extending
past posterior border of head; head widened
anteriorly and strongly narrowed posterior-
ly; vertex convex; maxillary and _ labial
palps, mesosoma, petiole and gaster similar
to that of C. longipilis.
Hairs erect, coarse and scattered on head,
mesosoma, petiole and gaster; decumbent
pubescence short and dilute on most sur-
faces.
Sculpture more coarse than other species,
head strongly and densely punctuate, in-
cluding surface of clypeus; mesosoma and
gaster with much weaker sculpture than
head, similar to other species except for C.
mirabilis.
Color medium to dark brown, head, man-
dibles and scapes dark brown, gaster with
slightly lighter splotches in same positions
as in C. mirabilis.
Type series.—Holotype 2, Pachitea,
Peru, Stdg; sphenocephalus Emery (MCSN)
[seen].
Material examined.—7 @, including
PERU: Pachitea (1 2 MCSN); Madre de
Dios, Cuzco Amazonico, 15 KNE Puerto
Maldonado, S. Cover & J. Tobin, vi-1989
(1 2 MCZC), Rio Tambopata Reserve, 30
air km SW Pto. Maldonado, 290m, 6/20-x1-
1979, J. B. Heppner (5 2 USNM).
Distribution.—Peru.
Discussion.—This species can be easily
separated from all others as the head, in-
cluding the clypeus, is strongly sculptured.
The head is strongly narrowed posteriorly,
similar to that found in C. leptocephalus. It
would not be confused with C. leptoceph-
alus, which has abundant erect hairs on the
top and bottom of the head, C. sphenoceph-
alus has only a few, scattered hairs on these
surfaces. This species is related to C. lon-
gipilis, but can be easily separated by the
sculpture of the clypeus and the shape of
the head.
203
Biology.— Unknown, collected in moist
tropical forest at Rio Tambopata Reserve.
ACKNOWLEDGMENTS
I thank the following curators and insti-
tutions for the loan of specimens: Mark
Smethurst (AMNH), Darrell Ubick
(CASC), Roy Snelling (LACM), Valter
Raineri (MCSN), Stefan Cover (MCZC),
David Smith (USNM). Emma MacKay,
Stefan Cover and an anonymous reviewer
provided a number of suggestions which
improved the manuscript. A number of in-
dividuals collected the specimens, especial-
ly S. Cover, J. Heppner, W. Mann, L. Pena,
W. Pulawski, E. Ross and J. Tobin. Without
their selfless efforts, ant systematics would
be in a worse state than it is.
This paper is dedicated to Dr. Valter Rai-
neri, curator of the Emery collection, with-
out whose help this revision and most of
the other taxonomic work I have done with
ants would not have been possible.
The research was supported by NSF
grant # HRD 9253021.
LITERATURE CITED
Emery, C. 1903. Intorno ad alcune specie di Cam-
ponotus dell’ America meridionale. Rendiconto
delle Sessioni della R. Accademia dell Scienze de-
Il’ Istituto di Bologna (N.S.) 7: 62-81.
. 1911. Fragments myrmécologiques. I-V. An-
nales de la Société Entomologique de Belgique
55: 213-225.
. 1920. Le genre ““Camponotus” Mayr. Nouvel
essai de sa subdivision en sous-genres. Revue
Zoologique Africaine 8: 229-260.
1923. Einige exotische Ameisen des
Deutschen Entomologischen Institutes. Entomo-
logische Mitteilungen 12: 60—62.
. 1925. Hymenoptera, Fam. Formicidae, sub-
fam. Formicinae. Jn Wytsman, P., ed., Genera In-
sectorum, Fasc. 183, 302 pp. Bruxelles.
Forel, A. 1914. Le genre Camponotus Mayr et les
genres voisins. Revue Suisse de Zoologie 22:
257-276.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 204
NOTE
Brachycerocoris camelus Costa (Hemiptera: Pentatomidae: Podopinae) in
India: New Country Record
In a recent paper on the Asian species of
Brachycerocoris Stal, the distribution of B.
camelus Costa was given as Sri Lanka and
China (Schaefer, O’Donnell, and Patton
1996, Oriental Insects 30: 203). I have now
found in my own collection a specimen
from Coimbatore, Madras State, India,
which is 250-300 miles (about 50 of them
water) from Sri Lanka. The specimen was
collected October 16, 1953, and remains in
my collection. It closely resembles the
specimens of B. camelus described from
China in Schaefer et al. (op. cit.), differing
only in several features. The matted setae
of the Chinese specimens are rubbed off on
part of the abdominal sterna of the Indian
specimen, revealing the deep reddish-brown
of the cuticle itself. The following tubercles
are broader, lower, and more rounded in the
Indian than the Chinese specimens: medial
tubercle on pronotum (Indian: tubercle
length 11% of body length, height 67% of
tubercle length; Chinese: 7% of body
length, height equal to length), and the me-
dial tubercle on the scutellum’s posterior
(Indian: tubercle length 13% of body
length, height 43% of tubercle length; Chi-
nese: 11% of body length, height 67% of
length); the tubercle between the eyes is
slightly more rounded in the Indian speci-
mens than in the Chinese specimens.
This specimen represents a new country
record, India.
Carl W. Schaefer, Department of Ecology
and Evolutionary Biology, University of
Connecticut, U-43, Storrs, CT 06269-3043,
US:As
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, pp. 205-207
Book REVIEW
Thrips Biology and Management. Edited
by Bruce L. Parker, Margaret Skinner, and
Trevor Lewis. NATO ASI Series A: Life
Sciences Vol. 276. Plenum Press, New
York, N.Y. 1995. 636 pp. $145.00 (cloth).
Thysanoptera, like other “‘minor’’ insect
orders, have been essentially neglected by
many entomologists in the past. Recently,
however, interest in the group has suddenly
begun to soar. This has been due at least in
part to the ever-increasing sophistication of
our agricultural and silvicultural systems
and their heightened emphasis on thrips,
mites, and other minute forms. Thysanop-
tera not only serve as vectors of phytopath-
ogens, but their ever-increasing resistance
to pesticides presents enormous manage-
ment concerns. The most important thrips
pests are confined to the family Thripidae
and include Frankliniella occidentalis (Per-
gande) [Western flower thrips, WFT],
Thrips tabaci Lindeman, and Thrips palmi
Karny.
In recognition of the upsurge in world-
wide interest concerning Thysanoptera, an
international conference was held in Bur-
lington, VT, U.S.A., in September, 1993,
entitled: ‘““Thysanoptera: Toward Under-
standing Thrips Management.” The vol-
ume, Thrips Biology and Management re-
sulted from this conference. The conference
organizers included, besides the listed vol-
ume editors, Bruce S. Heming, Carl C.
Childers, and Michael P. Parrella. The goals
of the organizing committee in convening
the conference as well as in production of
the text included: pointing out critical re-
search needs, obtaining fresh perspectives
on various biological and pathogen vector-
ing parameters and on the complex man-
agement issues, and finally, providing im-
portant background information for deci-
sion-making in agricultural/silvicultural
production, marketing, and quarantine sys-
tems. The conference hosted over 180 par-
ticipants representing over 25 nations. The
volume is a complete record of all oral (in-
vited) presentations, and most of the graph-
ic display presentations as well.
Following the Preface, the subject matter
is divided into the following 9 Sections:
Ife Introduction to Thrips/Plant Relation-
ships
II. | Pest Problems in Field, Forest and
Glasshouse Crops
III. Vectoring of Plant Pathogens
IV. Biological Control Agents and Prac-
tices
V. Chemical Control
VI. Integrated Pest Management
Biology and Behavior
Development and Genetics
IX. Field Sampling and Laboratory Tech-
niques
Section One (4 papers) begins with ex-
tensive review articles by Laurence A.
Mound and others concerning general
thrips biology with emphasis on feeding be-
havior. It was stressed that the opportunistic
feeding behaviors prevalent among Thysa-
noptera present a great potential for many
previously non-economic species to be-
come significant pests. The remainder of
this section concerns the effects of thrips
feeding and ovipositional activities on plant
host tissues, and on whole plants.
Section Two (14 entries) begins with an
excellent review by Thomas Leigh on the
pest thrips of cotton on a worldwide basis.
Following this is a series of articles on for-
est and orchard thrips pests in the U.S.A.
and New Zealand, most involving the pear
thrips, Taeniothrips inconsequens (Uzel).
Other inclusions in this section involve
thrips pests of vegetables and other crops
around the world, ending with a paper by
G. (Bert) Vierbergen on the international
movement, detection, and quarantine of
Thysanoptera pests.
206
Section Three (12 papers) consists main-
ly of articles concerning field and labora-
tory/electron microscopy studies of Tomato
Spotted Wilt Virus (TSWV) and its thrips
vectors, particularly F. occidentalis. Anoth-
er virus vectored by 7. palmi was reported
from India. This section closes with two pa-
pers on thrips vectoring/assisting the spread
of fungal phytopathogens. The Tospovirus-
es, including TSWV among others, are
transmitted by at least 8 species of thrips to
a wide variety of plant hosts, resulting in
serious epidemics in many economic crops
worldwide. Although in 1993, research on
TSWV and its relationships with its thrips
vectors at the tissue and cellular level was
only in its initial phases, it was already be-
ing shown that thrips could indeed serve as
primary vectors of the virus.
Section Four (21 entries) is primarily a
compilation of preliminary/basic research
reports on various aspects of thrips biolog-
ical control. Many articles involved Orius
(Heteroptera: Anthocoridae), and/or preda-
tory mites (Phytoseidae: Amblyseius). No-
table among these is one report from a Dan-
ish glasshouse nursery concerning their
**Keep Down” approach. Inundative releas-
es of Orius onto potted ornamentals when
thrips were at initial low levels (BEFORE
thrips reached pre-determined economic
threshold levels), and maintaining thrips
populations below economic levels resulted
in high quality plants for the market. Ad-
ditional entries in this section include a pa-
per that discusses nematode parasitism of
Thysanoptera, two papers on fungal patho-
gens of thrips (one, an extensive literature
review) and finally, three articles emphasiz-
ing Thysanoptera as biological control
agents for other economic pest groups, usu-
ally spider mites (Tetranychidae). Notable
among the latter is a review article on the
situation in cotton in the San juaquin Valley
of California. Here, F. occidentalis has
come to be considered a beneficial species
which feeds on three species of spider
mites. When mites are at high population
levels, WEFT serves as an important mite
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
predator, but when mite populations are
low, WEFT then assumes importance as a
prey species for a complex of generalist
predators. These predators and WFT keep
the mites under excessive levels at times
when mites would otherwise become prob-
lematic. This has resulted in markedly re-
duced needs for chemical controls in San
Juaquin Valley cotton.
Section Five (3 inclusions) titled ““Chem-
ical Control” was primarily concerned with
resistance studies and the limitations of cer-
tain fumigation treatments. The first two ar-
ticles concern resistance studies using di-
verse populations of WEFT exposed to var-
ious pesticides. The last paper reports on a
study of the fumigants naled and sulfotep,
prospective replacements for methyl bro-
mide, and the failure of either agent to
achieve regulatory control of WEFT in cut
flower commodities (Dianthus).
Section Six (12 entries) contains 7 papers
on WFT integrated/non-chemical manage-
ment approaches, with the remaining papers
in this section involving other species of
thrips. As is true with the majority of the
sections in this book, there is a mixture of
review articles and basic/field research re-
ports as well as other miscellaneous papers.
There is an excellent “how-to” article
which a nurseryman could use in designing
an IPM program for greenhouse thrips man-
agement. Several papers are concerned with
host plant resistance to WEFT and other
thrips; results with both flower crops and
vegetables (cucumber) are promising.
Section Seven (11 papers) begins with an
important article which summarizes over a
decade of research in and around New York
vegetable fields, concerning primarily T. ta-
baci and its relationships to cabbage. An-
other paper concerns the results of over 6
years of research in Georgia which eluci-
dated population trends in the various thrips
involved, and their inter-relationships with
TSWYV levels in peanut fields. Another im-
portant long-term study explores population
trends in thrips of peach and nectarine or-
chards in Pennsylvania. A final long-term
VOLUME 99, NUMBER 1
study concerns Thysanoptera species in
meadows of the southern Carpathian moun-
tains of Romania, and delineates relation-
ships between species richness and altitude.
Finally, two exquisite behavioral studies are
included. One involves laboratory exami-
nation of chemical communication between
WFT larvae and their predators, and the
other examines detailed field behavior of
natural male WFT aggregations in Utah.
Section Eight (3 submissions) most no-
tably includes Gerald Moritz’s evidence
(using electron microscopy) that Thysanop-
tera are indeed Holometamorphic, and ter-
minology for developmental stages in
Thysanoptera should be egg, larva, pupa,
adult.
Section Nine (11 entries) stresses meth-
odology, and concerns survey techniques
for field studies, as well as laboratory rear-
ing/processing techniques.
In summary, Thrips Biology and Man-
agement is a volume that contains the work
of thrips workers worldwide, and much of
the included material involves basic re-
207
search on F. occidentalis. It is well indexed,
with: an index to thrips species/synonyms
mentioned in the text; a list of conference
participants; a thrips species index (those
referred to in the text); and, finally, a sub-
ject index. Typographical errors were non-
problematical other than the one on page
376, at the end of the first line of the last
paragraph on the page: “‘in the early
1990’s”’ should read “‘in the early 1900's.”
Finally, persons who would find this
book to be of interest and invaluable, in-
clude not only economic entomologists, but
also general entomologists, federal and state
policy makers, agricultural quarantine and
extension personnel, commercial nursery-
men (including those in greenhouse opera-
tions), farmers, crop scouting specialists,
and anyone with a broad interest in various
aspects of agriculture and forest pest man-
agement.
Susan Broda-Hydorn, U.S.D.A., A.P.H.LS.,
P.P.Q., 2200 Broening Hwy., Suite #140,
Baltimore, MD 21224, U.S.A.
PROC. ENTOMOL. SOC. WASH.
99(1), 1997, p. 208
OFFICIAL SEAL OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
The following history of the official seal
appeared in the 1964 issue of the Proceed-
ings of the Entomological Society of Wash-
ington (66(1): 1) and was written by Jon L.
Herring, editor at that time.
The Official Seal of the Entomological
Society of Washington
The story of the discovery of Rheuma-
tobates rileyi Bergroth and its subsequent
portrayal on the cover of our proceedings
is an interesting one.
This small water strider was first men-
tioned in 1891 by Reverend J. L. Zabriskie
(Jour. N. Y. Micro. Soc. 7: 128-129, 1 fig.)
in an article entitled ““A curious, unknown,
aquatic, hemipterous larva.” Then, in 1892,
there appeared anonymously (Insect Life 4:
198) a complete description and excellent
illustration of this bug, but it remained un-
named. On the basis of this description,
Bergroth proposed the name Rheumato-
bates rileyi, new genus and species without
further description (Insect Life 4:321). All
of the above references dealt with apterous
forms.
Otto Heidemann, specialist in Hemiptera,
with the old Bureau of Entomology, col-
lected the first winged specimens near Glen
Echo, Maryland in 1893. Mr. Heidemann,
who, in addition to his other talents, was a
gifted engraver, made a handsome woodcut
of a fully winged male. This figure ap-
peared for the first time on the cover of vol-
ume 3, number | of our Proceedings (issued
March 8, 1894).
The cut appeared continuously through
1920. It is mentioned in Mr. Heidemann’s
obituary in 1916 (Proc. Ent. Soc. Wash. 18:
202) that the illustration had been adopted
as the official seal of our society. However,
I am unable to verify this from the minutes
of these early meetings.
The seal was dropped from the cover in
1921, presumably because the woodblock
needed replacing. Later a photograph of the
seal was made and a new illustration ap-
peared on the June 1937 issue and has ap-
peared continuously since then. Because
much detail in the original has been lost, I
have redrawn the seal, based on specimens
from Glen Echo, and had a new cut made.
The results can be seen on the cover of this
issue.
The seal is widely recognized as a sym-
bol of our society. As an example, one of
the late Professor H. B. Hungerford’s exam
questions was, “‘Name the society whose
official seal is Rheumatobates rileyi?”
Editor [Jon L. Herring]
PUBLICATIONS FOR SALE BY THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON
MISCELLANEOUS PUBLICATIONS
Cynipid Galls of the Eastern United States, by Lewis H. Weld. 124 pp. 1959 $ 5.00
Symp alls of the Southwest, by Lewis H. Weld. 35 pp. 1960 3.00
Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C.
LTS SEILS BIST Oy aR US IPA 7 eet BE AAS RAR ToS SRA Ie eR ee 1.50
Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C.
SUSI. Gilroy ove SIT oo MR seh SS IONE ee en ee ee ee 2.00
A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael
1S SICUDELBL A ohS 5 0) opat I 10 een a PE RIS a eS Se BR ee Ey eh ee aa 10.00
MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 __. $15.00
No.2. A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving. (out of
WSS) Yayo). US Yar ee ob ei ett abe Able es eee Ce ern DNS a eo Oe Re cleranE Ceseeeeer ge we aU OR print)
No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman.
PSS BVO AICS) ae eR ROR LS ipa ie EER eS SI et LS eb 15.00
No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952 15.00
No.5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 _- 15.00
No.6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi
AKAN ASI. 2 SON PP al OO eos eas teed Vee B As ae ae ee ees ele Veh Many ae ease a eet Vee 15.00
No.7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp.
IED TG pe tesla oe ci eas Sate CT Reis Son ON ee ee 2 Se Re SSS ER Bh 11.00
No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato-
porzonidas), by W. 1. Grogan. Jr and: W. W. Wirth: 125)pp: 1979" 22 ee 12.00
No.9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson.
EURO PYG ae LS) $y Lipps ie Ws LANE Ws aR le ERC SI A OE Soe a Bo Ge 0s Ly Wa ems COE Be YS AL 10.00
No. 10. Recent Advances in Dipteran Systematics: Commemorative Volume in Honor of Curtis W.
Sabrosky. Edited by Wayne N. Mathis and F. Christian Thompson. 227 pp. 1982 ___. 11.00
No. 11. A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370
ja) = TIO LS AS), Meee ae AR Sees ein EEE SA ees RSPAS he Pate eae I A a NE, 18.00
No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff.
SVT BY OUST Ua EVER SEN 5.00
No. 13. An Identification Manual for the North American Genera of the Family Braconidae (Hyme-
noptera), by Paul M. Marsh, Scott R. Shaw, and Robert A. Wharton. 98 pp. 1987 __.._ 18.00
No. 14. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174
(BR, TNS DIY I es I a ae ao PRR SE ALN ge ND NNT ek ee 25.00
No. 15. A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent
BnGaWe Be | Gropal ral S! Ppl OO Dy ss 2 ae tS eh Ee ce oe ee Pe 25.00
No. 16. The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 __. 25.00
No. 17. Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Krombein, edited
Dyabe be Nordentand Azse, Menke .2116) pp 199 Gree eee ae ere een anes eee ee 25.00
No. 18. Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and
AMIE a oy) BIN Gs Rovere vy etc W nays PALIT a0) 0 el We 2 0 fe esate renee mete als ceo ieee a UN ee) EI gle I ee ee 25.00
Back issues of the Proceedings of the Entomological Society of Washington are available at $60.00 per volume
to non-members and $25.00 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, % Department of Entomology, MRC 168, Smithsonian
Institution, Washington, D.C. 20560.
CONTENTS
(Continued from front cover)
MATHIS, WAYNE N.—A review of the shore-fly genus Diphuia Cresson (Diptera: Ephy-
dridae)
MEJDALANI, GABRIEL and MARCIO FELIX—A new species of the Neotropical genus Lis-
soscarta Stal (Homoptera: Cicadellidae: Cicadellinae) that mimics wasps
MENKE, ARNOLD S.—A new Ammoplanus Gussakovskij (Hymenoptera: Sphecidae: Pem-
phredoninae) from San Clemente Island, California
MUNOZ-QUESADA, FERNANDO—Five new species and a new record of Costa Rican Lep-
tonema Guérin (Trichoptera: Hydropsychidae)
PIKE, K. S., P. STARY, D. ALLISON, G. GRAF, L. BOYDSTON, and T. MILLER—Parasit-
oids (Hymenoptera: Braconidae: Aphidiinae) of aphids on big sagebrush (Artemisia triden-
tata Nuttall) and prairie sage (Artemisia ludoviciana Nuttall) in Washington State
ROSE, MIKE and GREGORY ZOLNEROWICH—Eretmocerus Haldeman (Hymenoptera:
Aphelinidae) in the United States, with descriptions of new species attacking Bemisia (tabaci
complex) (Homoptera: Aleyrodidae)
SCHAEFER, CARL W. and JOSEPH C. SCHAFFNER—The New World alydine genus Stach-
yocnemus Stal (Hemiptera: Alydidae)
SLATER, JAMES A. and HARRY BRAILOVSKY—A new species of Drymus Fieber from
Mexico, with a key to species and a checklist of Western Hemisphere Drymini (Hemiptera:
Lygaeidae)
SMITH, DAVID R.—Aulacidae (Hymenoptera) of Sri Lanka
WIJESEKARA, G. A. W. and M. E. SCHAUFF—Two new genera and three new species of
Euplectrini (Hymenoptera: Eulophidae) from the New World
NOTE
SCHAEFER, CARL W.—Brachycerocoris camelus Costa (Hemiptera: Pentatomidae: Podo-
pinae) in India: new country record
BOOK REVIEW
BRODA-HYDORN, SUSAN—Thrips Biology and Management, edited by Bruce L. Parker,
Margaret Skinner, and Trevor Lewis
MISCELLANEOUS
Official seal of the Entomological Society of Washington
VOL.99 4G! APRIL 1997 NO. 2
Eo | ed (ISSN 0013-8797)
oe eae
PROCEEDINGS
of the
ENTOMOLOGICAL SOCIETY
CONTENTS
ADAMSKI, DAVID and BERNARD LANDRY—Review of the Blastobasinae (Lepidoptera:
Gelechioidea: Coleophoridae) of the Galapagos Islands ................... 2.2 cece cece eeeee eee 348
BRAILOVSKY, HARRY and ERNESTO BARRERA—Redescription of the subgenus Hygia
(Eucolpura) Breddin (Hemiptera: Coreidae: Colpurini), with the description of two new
Species, anda! key sto) the) Known) SPECIES 20/15 tea alswere cteicte ee s)oisls Sele «je areintt nioialncidinie olalfolelespjsie)~ oicieloh 257
BUENO-SORIA, JOAQUIN and SILVIA SANTIAGO-FRAGOSO-—Studies in aquatic insects
XII: descriptions of nineteen new species of the genus Ochrotrichia Mosely (Trichoptera:
Hydroptilidae) from Mexico and Central America .......0...........0.cee bec ctee eset ee eee ees 359
FITZGERALD, SCOTT J.—Additional notes on Nearctic Bibio Geoffroy (Diptera: Bibion-
Wie) ERS aie iy a Ae eee ard, Raa aE CEOS EING APES bE Bama Seo Seog dation Mle ob op rtie esq anb cic: 294
GAGNE, RAYMOND J., JOSEPH K. BALCIUNAS, and DAMIEN W. BURROWS—Six
new species of gall midges (Diptera: Cecidomyiidae) from Melaleuca (Murtaceae) in
PNT SISTER NA Rie es AD Mine Aaah Sn Re anu yen AL Nels Ue RA AW Abert ina 48 312
GANDOLFO, DANIEL E. and ALLEN L. NORRBOM—A new species of Trypanaresta Hering
(Diptera: Tephritidae) from Patagonia, a potential agent for biological control of snakeweeds
KGHIERREZIAUSPps) MME ME MINIT CA) SLATES WV: tele ah nalts as eats aafees nel Ae yaaa ae ay ainloith alate sleitpel alee icy iste sta 248
HODGES, RONALD W.—A new agonoxenine moth damaging Araucaria araucana needles in
western Argentina and notes on the Neotropical agonoxenine fauna (Lepidoptera: Gelechioi-
EE LACIISTI ACI nat nated Pe ache ei hace ea ca sien b cleoays ire Se yan MoO GPRS el ae oP uals ae camp cists ch staja Ghat 267
KIRCHNER, RALPH F. and BORIS C. KONDRATIEFF—A new species of Nearctic Perlesta
(Plcasopteradberidac) conn Nil einid: hi. c+ one c teerae oes stitial ese eee eer oe ele aelnla etelors 290
MARSHALL, S. A.—Limomyza, a new genus of primitive Limosininae (Diptera: Sphaeroceri-
dae), with five new species from United States, Mexico, and Central America .............. PAS)
MILLER, DOUGLASS R. and DOUGLAS J. WILLIAMS—A new species of mealybug in the
genus Pseudococcus (Homoptera: Pseudococcidae) of quarantine importance ................ 305
NORRBOM, ALLEN L.—The genus Carpomya Costa (Diptera: Tephritidae): new synonymy,
description of first American species, and phylogenetic analysis ............-..2..0-.eeeeee ees 338
(Continued on back cover)
x A
THE
ENTOMOLOGICAL SOCIETY
OF WASHINGTON
ORGANIZED MARCH 12, 1884
OFFICERS FOR 1997
M. ALMA SOLIS, President MICHAEL G. POGUE, Treasurer
WARREN E. STEINER, JR., President-Elect DaAviD G. FURTH, Program Chair
HAROLD HARLAN, Recording Secretary DaviD ADAMSKI, Membership Chair
HOLLis B. WILLIAMS, Corresponding Secretary RALPH P. ECKERLIN, Past President
Gary L. MILLER, Custodian
DaviD R. SmitH, Editor
Publications Committee
THOMAS J. HENRY WAYNE N. MATHIS
GARY L. MILLER, Book Review Editor
Honorary President
Curtis W. SABROSKY
Honorary Members
LouIsE M. RUSSELL ALAN STONE KARL V. KROMBEIN
All correspondence concerning Society business should be mailed to the appropriate officer at the following
address: Entomological Society of Washington, % Department of Entomology, MRC-168, Smithsonian Insti-
tution, Washington, D.C. 20560.
MEETINGS.—Regular meetings of the Society are held in the Natural History Building, Smithsonian Institu-
tion, on the first Thursday of each month from October to June, inclusive, at 8 P.M. Minutes of meetings are
published regularly in the Proceedings.
MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology.
Annual dues for members are $25.00 (U.S. currency).
PROCEEDINGS.—The Proceedings of the Entomological Society of Washington (ISSN 0013-8797) are pub-
lished quarterly beginning in January by The Entomological Society of Washington. POSTMASTER: Send
address changes to the Entomological Society of Washington, % Department of Entomology, MRC-168, Smith-
sonian Institution, Washington, D.C. 20560. Members in good standing receive the Proceedings of the Ento-
mological Society of Washington. Nonmember U.S. subscriptions are $60.00 per year and foreign subscriptions
are $70.00 per year, payable (U.S. currency) in advance. Foreign delivery cannot be guaranteed. All remittances
should be made payable to The Entomological Society of Washington.
The Society does not exchange its publications for those of other societies.
PLEASE SEE PP. 615-616 OF THE JULY 1996 ISSUE FOR INFORMATION 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, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash-
ington, D.C. 20560.
Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology,
Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560.
Books for Review: Gary L. Miller, Systematic Entomology Laboratory, ARS, USDA, Building 046, BARC-
West, Beltsville, MD 20705.
Managing Editor and Known Bondholders or other Security Holders: none.
This issue was mailed 15 May 1997
Second Class Postage Paid at Washington, D.C. and additional mailing office.
PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA
This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 209-237
REVISION OF THE ERISTALIS FLOWER FLIES (DIPTERA: SYRPHIDAE) OF
THE AMERICAS SOUTH OF THE UNITED STATES
E CHRISTIAN THOMPSON
Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart-
ment of Agriculture, c/o National Museum of Natural History, MRC 168, Washington,
DC 20560, U.S.A.
Abstract.—Three new species are described (Eristalis alleni, E. gatesi, and Palpada
eristaloides); lectotypes are designated for bogotensis Macquart, circe Williston, latifrons
Loew, rufoscutata Bigot, stipator Osten Sacken; new synonyms noted (Eristalis tenuifrons
Curran 1930 = Palpada semicircula Walker 1852; Eristalis aztecus Hull 1935 = circe
Williston 1891; Eristalis colombica Macquart 1855 = tenax Linnaeus 1758); and all
species redescribed (Eristalis bellardi Jaennicke and persa Williston).
Key Words: key, neotropical
The Biodiversity Crisis is now the rage
(NBS 1989; Wilson 1985a, 1988, 1992).
Much has been written in both the scientific
and popular literature. The world biota is
disappearing at an alarming rate never be-
fore equaled in the history of this universe.
That is not disputed. What we do not really
know is what is being lost as our knowl-
edge of our biota is abysmal. The big crea-
tures, such as birds and mammals, as well
as flowering plants, are known, named and
classified, but the little ones, from the small,
such as flies, worms and fungi, to the mi-
croscopic, such as protozoans, bacteria and
Viruses, are not well known, mostly unna-
med and poorly classified (Erwin 1992;
May 1990, 1992; Wilson 1987). To make
them known will require the collaborative
effort of many people, from collectors and
taxonomists to computer scientists, states-
men and philanthropists (Raven and Wilson
1992; Roberts 1988; Yoon 1993). Costa
Rica has started to build this collaboration.
Their view of conservation is that biodi-
versity can only be conserved if people
deem biodiversity to be of value. So, con-
servation is a process of saving it, then
knowing it, so that it can be used sustain-
ably (Janzen 1991). Costa Ricans have set
aside a significant portion of their land to
conserve wild biodiversity. They have es-
tablished a national institute for biodiversity
(INBio) to develop a cadre of professionals
to inventory Costa Rican biodiversity and
then to help others to find sustainable and
profitable uses of this biotic wealth (Tan-
gley 1990, Hovore 1991, Gamez and Gauld
1993). This inventory requires a universal
naming system, which is provided by tax-
onomy (Thompson 1996). Taxonomy is
global science, as many organisms are wide
ranging and the groups that they are placed
in are not necessarily defined by geography.
Taxonomists, who tend to be clustered in
the developed North (Gaston and May
1992), are themselves a dying breed (Wil-
son 1985b, Holden 1989, Culotta 1992, No-
vacek 1992). So, for taxonomists, the chal-
lenge is not only to name and classify our
unknown biota before it is lost, but also to
do it before they themselves are extinct! Or
the alternative is to demonstrate more clear-
210
ly their value. So, together some taxono-
mists have begun to work with Costa Ri-
cans to inventory their biodiversity. This
paper, which describes two new and spec-
tacular flies known only from Costa Rica,
is one result of this unique collaboration.
The pair of new species herewith de-
scribed are most striking and beautiful spe-
cies of Eristalis (Figs. 1, 2) that I know.
Hence, I have named them after the brilliant
pair of Paul Allen and Bill Gates, who have
done more to bring power to the people
through computer software. That in turn has
allow for the blossoming of all of our tal-
ents, not the least the ability of systematists
to more effectively and efficiently describe
our disappearing biota.
Eristalis flower flies are common, some-
times abundant, in northern and montane
temperate habitats. The adults are pollina-
tors and the larvae, called rat-tailed mag-
gots, are filter-feeders in organically rich
waters, such as ponds, pools and ditches,
aiding in nutrient recycling. Because of
their diverse life-cycle, the species are good
indicators of the health of their environ-
ments.
Eristalis was a broadly defined group
when first established by Latreille (1804)
and that definition was further enlarged by
Fabricius (1805), whose interpretation was
characterized as ‘“‘ein mischmasch”’ by Mei-
gen (1822: 382). Meigen restricted the
name to those species with sinuate vein
R4+5 and a petiolate cell R1 (although he
didn’t use these characters). Except for Zet-
terstedt (1837, 1838, 1842—60), who incor-
rectly used the name for Cheilosia of au-
thors, all subsequent workers have followed
Meigen’s definition of Ervistalis. Later, some
authors, such as Rondani (1845, 1857) and
Mik (1897), did subdive Eristalis into
smaller components, but most authors con-
tinued to the use the name in the broad
sense. The first and only modern attempt at
subdivision of Eristalis was by Kanervo
(1938), but his work was ignored. Vocker-
oth (in litt.) and Thompson (1972) re-ex-
amined Kanervo’s work, and divided Eris-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
talis into a few monophyletic groups. Most
of the Neotropical species previously
placed in Ervistalis belong to Palpada; only
a few belong to Eristalis, sensu stricto. A
key to separate the Neotropical components
of Eristalis of prior authors is appended.
There has never been a revision of the Neo-
tropical species of Eristalis (sensu lato).
However, Curran (1930, 1934) did publish
keys, which covered most of the species.
The Eristalis flower flies of Latin Amer-
ica present an interesting zoogeographic
problem. Unfortunately, a character analy-
sis of the genus Eristalis as whole has not
been done, so a zoogeographic analysis of
the Latin American species is speculative.
What prior authors called E. bogotensis is
a superspecies, consisting of E. bellardii in
Mexico, E. bogotensis in northern and cen-
tral South America and E. croceimaculata
in southern South America. The E. bogo-
tensis superspecies is characterized by a
monomorphic abdominal pattern which
consists of large reddish to orange lateral
maculae, the extent of which varies clinally:
Eristalis bellardii has the most extensive
pale maculae, whereas E. croceimaculata
has the least extensive pale maculae, and E.
bogotensis is intermediate. This clinal vari-
ation in color has been noted in other flies
and insects (see Zuska and Berg 1974) and
is related to temperature. There is no com-
ponent of the E. bogotensis superspecies in
Central America. In Middle America, there
are 4 species. In northern areas (Mexico
(Chiapas) and El Salvador), there is a pair
of sympatric sexually dimorphic species, E.
circe and E. persa, in which the male has
an orange abdomen and the female has a
black abdomen. In the south (Costa Rica),
there are two sympatric monomorphic spe-
cies, one with an orange abdomen and the
other with a black and yellow abdomen.
The scenario these distributions and char-
acters suggest is that there was a wide-
spread ancestral species, which vicariated
into 4 allopatric components, with the Mid-
dle American component undergoing fur-
ther vicariance [Ancestral species = E. bel-
VOLUME 99, NUMBER 2 211
2
SF 1993
© Thompson 1993
Figs. 1-2. Adult, dorsal view. 1, Eristalis alleni. 2, E. gatesi. ©Thompson 1993.
lardii + ((E. circe + E. persa) + (E. gatesi America have richer and more diverse fau-
+ E. alleni)) + E. bogotensis + E. crocei- nas that those to the north or south, which
maculata]. The significance of such a sce- is not unusual given the isolated nature of
nario is that the higher elevations of Middle these areas (Fig. 3). Unfortunately, these ar-
212 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Goh
‘ 7 -
Ne
Fig. 3. Middle America showing in black the areas of 2,000 meters or higher.
eas are limited in size and threatened by
development.
KEY TO GENERA OF ERISTALIS-LIKE FLOWER
FLIES FOUND IN THE NEW WORLD
1. Postalar pile tuft present; eye maculate; ane-
pimeron with dorsomedial and posterior por-
tions pilose; katepimeron bare; arista bare . . .
Os ee eens CRISES Soe oye Eristalinus
— Postalar pile tuft absent; eye plain, without
dark maculae; anepimeron with dorsomedial
and sposterlon portions) bate) eae ee 2
. Meron with pile anterior to and/or ventral of
posterior spiracle; katepimeron pilose. Eye
with uniform pile; wing microtrichose or bare
Oe AN Eo acco GLO eae ae ene net oe Palpada
— Meron without pile near posterior spiracle;
Win Palbane™ erty cua eect ESVUSTQLIS err tee ee 3
3. Katepimeron pilose; eye pilose, with 2 vertical
tO
vittae of darker, contrasting pile; arista pilose
basally oo... yn oe Bierce hee Eristalis (s.s.)
— Katepimeron bare; eye uniformly pilose, with-
out darker vittae; arista bare or pilose ......
Fos aI >. 2 Oe SG eee! 8 hes ae E. (Eoseristalis)
KEY TO ERISTALIS-LIKE FLOWER FLIES OF
AMERICAS SOUTH OF THE UNITED STATES
= KeatepimeronipllOoSe yen itn ee 3
Katepimeron bates) 5s eee D
2. Meron without pile anterior to or ventrad of
spiracle; arista with short but distinct pile on
basal %; eye densely pilose, with 2 vertical
vittae of darker pile; wing bare .......
Meron with pile anterior to and/or ventrad of
spiracle; arista bare; eye pilose dorsally, with-
out contrasting fascia of pile; wing microtri-
chosejor bate 5 wee ee Palpada species
VOLUME 99, NUMBER 2
. Arista bare
Arista pilose at least on basal half
. Legs entirely black; pleura and femora black
pilose; frons extensively yellowish-white pi-
fosen(GostawRica)) 9. see Eristalis gatesi
Legs partially orange, at least on basal % of
tibiae; pleura orange to yellowish pilose; fem-
oral and frontal pile variable, usually femora
partially yellow pilose and frons with at least
some black pile
. Hind tarsus black; fore and mid tibiae with
apical %4 or more black; fore and mid tarsi
brownish black apically; mesonotum includ-
ing postalar callus and scutellum entirely or-
ange to fulvous pilose; male abdomen orange
with medial black maculae; female abdomen
black (Mexico) Eristalis circe
Hind tarsus, fore and mid tibiae, and tarsi or-
ange; postalar callus at least partially black
pilose; mesonotum and scutellum frequently
partially black pilose
Hind tibia almost entirely black, only with
base and apex narrowly orange; face black;
female abdomen entirely black; male abdo-
men orange to fulvous pilose; male fore fe-
mur with basoposterior tuft of long dense
black pile (Mexico to El Salvador)
Eristalis persa
Hind tibia extensively orange, rarely with in-
distinct medial brownish annulus; face exten-
sively tawny to orange in ground color; fe-
male abdomen reddish orange with medial
black maculae; abdomen extensively black pi-
lose; male fore femur without such a tuft of
long pile (Costa Rica)
Eristalis alleni
. Mesonotum with 3 transverse gray pollinose
fasciae (Costa Rica, Panama)
asc: Ct EAA ARA, Steer eas Palpada semicircula
Mesonotum without pollinose fasciae
. Second tergum with a complete posterior
black fascia (northern Mexico)
rts Lg ete Sot See Eristalis stipator
Second tergum entirely reddish laterally,
without a complete posterior black fascia,
with black color restricted to medial area .. 9
. Antenna orange; hind tibia entirely orange;
face orange (Ecuador) .... Palpada eristaloides
Antenna dark, brown to black; hind tibia
black on apical % or more; face black medi-
ally
. Coxae, katepisternum and metasternum yel-
low pilose; male with orange abdominal color
extending laterally to apex of 4th tergum;
front of female usually entirely pale pilose on
ventral *4; male eye contiguity much longer
than vertical triangle (2.5 to 1.5 times) (Mex-
ico) Eristalis bellardii
Coxae, metasternum and katepisternum ven-
trally partially to entirely black pilose; male
with orange abdominal color extending only
to base of 4th tergum; front of female par-
tially black pilose on ventral *%4; male eye con-
tiguity shorter than vertical triangle (0.9
times) (South America)
11. Female: 3rd tergum completely black; 2nd
tergum with sides narrowly dark and posterior
margin black on medial % or more. Male: 2nd
tergum black on posteromedial % or more; 4th
tergum completely black; male genitalia usu-
ally black pilose; aedeagus with ventral
curved prongs narrow; superior lobe with
only a short lateral carina (Chile, southern Ar-
gentina)
— Female: 3rd tergum orange basolaterally; 2nd
tergum with sides entirely orange, with pos-
terior margin black only on medial % or less.
Male: 2nd tergum with posterior margin black
on medial % or less; 4th tergum frequently
narrowly orange basolaterally; male genitalia
yellow pilose; aedeagus with ventral prongs
broad, flared; superior lobe with long carina
extending from base to apex (Columbia, s. to
Peru, Bolivia, & northern Argentina)
BS ee Re ee ee Eristalis bogotensis
Eristalis croceimaculata
Genus Eristalis Latreille
Tubifera Meigen 1800: 34. Type species,
Musca tenax Linnaeus (subsequent des-
ignation by Coquillett 1910: 618). Sup-
pressed by ICZN 1963: 339.
Elophilus Meigen 1803: 274. Type species,
Musca tenax Linnaeus (subsequent des-
ignation by Latreille 1810: 443). Sup-
pressed by ICZN 1993; 256.
Eristalis Latreille 1804: 194. Type species,
Musca tenax Linnaeus (subsequent des-
ignation by Curtis 1832: pl. 432).
Eristaloides Rondani 1845: 453. Type spe-
cies, Musca tenax Linnaeus (subsequent
designation of Coquillett 1910: 540).
Eristalomya Rondani 1857: 38. Type spe-
cies, Musca tenax Linnaeus (original des-
ignation).
Eriops Lioy 1864: 743 (preocc. Klug 1808).
Type species, Musca tenax Linnaeus
(subsequent designation by Goffe 1946:
29):
Head: Face broadly pilose and pollinose
laterally, usually shiny and bare medially,
rarely entirely pollinose, straight except for
medial tubercle and slight anterior produc-
214 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tion at antennal pits; tentorial pit short, ex-
tending along ventral third of eye; facial
stripes indistinct; frontal prominence low,
on dorsal third of head; eye pilose, holoptic
in males; antenna short, about % as long as
face; basoflagellomere quadrate, slightly
longer than broad.
Thorax: Slightly longer than broad, with
long pile; anterior anepisternum bare; ka-
tepisternum continuously pilose from ven-
tral to dorsal margin; anepimeron with dor-
somedial and posterior portions bare; me-
tathoracic pleuron bare; metathoracic spi-
racle, large, larger than basoflagellomere;
plumula long and multibranched.
Legs: Simple, hind femur not swollen,
hind tibia without basal nor apical carina
nor apical dens. Wing: Bare; cell R1 closed
and petiolate; cell R4+5 petiolate, with pet-
iole as long as stigmatic crossvein; stig-
matic crossvein present.
Abdomen: Oval to suboval.
Subgenus Eristalis
Eye with two denser vertical vittae of
pile; arista bare; katepimeron pilose.
The subgenus Eristalis contains only 2
species, E. proserpina Wiedemann from
China, and E. tenax Linnaeus, an Old World
synanthropic species which is now cosmo-
politan.
Eristalis (Eristalis) tenax (Linnaeus)
(Figs. 4, adult; 5, larva)
Musca tenax Linnaeus 1758: 591 Sweden
(restricted Thompson et al. 1982: 160).
LT 3 LSL (designated Thompson et al.
1982: 160).
Eristalis tenax: Bréthes 1907: 293 (Argen-
tina, catalog citation); Kertész 1910: 238
(catalog citation); Porter 1921: 447,
1924a: 82, 1924b: 98, 1927: 122, 1928:
224. 1932: 190, 1934: 170, 1938-155
(Chile); Hull 1925: 305 (description);
Curran 1930: 6, 1934: 410 (key refer-
ences); Shannon & Aubertin 1933: 163
(Chile); Ruiz & Stuardo 1935: 318
(Chile); Gutierrez 1939: 35 (Chile); En-
derlein 1940: 662 (Juan Fernandez Is.);
Stuardo 1946: 128 (catalog citation,
Chile); Fluke 1957: 144 (catalog citation,
Chile, Argentina); Etcheverry & Shene-
felt 1962: 208 (Male genitalia figured);
Etcheverry 1963: 44 (synonymy, Chile);
Thompson et al. 1976: 101 (catalog ci-
tation); Thompson 1972: 140 (Male gen-
italia figured), 1981: 146 (West Indies,
Status).
Eristalomyia tenax: Enderlein 1940: 662
(Juan Fernandez Islands).
Tubifera tenax: James 1947: 151 (habitus
illustrated, description, myiasis, distr.).
Eristalis columbica Macquart 1855: 108 @
Columbia. LT 2 BMNH here designated.
Williston 1886: 319 (catalog citation);
Kertész 1910: 215 (catalog citation);
Fluke 1957: 154 (catalog citation, sp. in-
certa sedis). New synonym.
Palpada columbica: Thompson et al. 1976:
104 (catalog citation)
Male.—Head: Black; face yellowish-
gray pollinose except broad shiny medial
vitta, yellow pilose; gena shiny, yellow pi-
lose; frontal lunule brownish orange except
black medially; frontal triangle shiny and
black pilose apicomedially, yellowish-gray
polllinose and yellow pilose laterally; ver-
tical triangle dull black pollinose, black pi-
lose; eyes holoptic, with eye contiquity
short, about as long as ocellar triangle; oc-
ciput yellowish-gray pollinose, yellow pi-
lose; antenna brownish black; arista bare,
brown.
Thorax: Black, yellowish gray pollinose,
yellow pilose, postalar callus yellow pilose;
scutellum yellow, shiny, yellow pilose;
Ssquama and plumula orange yellow; spirac-
ular fringes light brownish yellow; halter
yellow, with brown head; wing hyaline,
bare; epaulet yellow pilose; basicosta black
and yellow pilose. Legs: Brownish black
except orange femoral-tibial joints and mid
basotarsomere on basal %4, yellow pilose ex-
cept hind femur black pilose apicoventrally
on posterior edge, hind tibia black pilose
posteriorly and hind tarsus black pilose.
Abdomen: Brownish yellow pilose; Ist
VOLUME 99, NUMBER 2
Figs. 4—5.
sternum gray pollinose, brownish black ex-
cept paler laterally; 2nd sternum shiny, yel-
low except medial % brownish black; 3rd
sternum shiny, black except paler basola-
terally; 4th sternum shiny, black; Ist tergum
gray pollilnose, black; 2nd tergum black ex-
cept orange apical margin and large medio-
lateral ovoid orange macula, with macula
occupying full lateral width and extending
to medial 4; 3rd tergum shiny, black except
narrow orange apical, broader orange an-
terior margin except medial %, and broad
215
\ ee fs ‘ is
Pe . Cac
— Cu ~ eo i eos. 1 ae J (/ ‘
Se ae ee USA, —
os y
LPG:
Eristalis tenax. 4, Adult, dorsal view. 5, Larva, lateral view.
orange mediolateral fascia, with fascia con-
nected basolaterally to pale anterior margin,
with fascia from ¥, to % tergal length; 4th
tergum shiny, black, rarely narrowly orange
basolaterally; genitalia shiny, black.
Female.—Similar; front shiny dorsad lu-
nule and anterior to ocellar triangle, else-
where yellowish gray pollinose, which is
lighter ventrolaterally and darker dorsome-
dially, extensively black pilose, with yellow
pile intermixed medially and ventrolateral-
ly, about .4 times as wide as head at anten-
216 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
na, tapering to about .35 times as wide dor-
sally; frequently with abdominal orange
maculae reduced, sometimes greatly so, in
such specimens no pale macula on 3rd ter-
gum and macula on 2nd tergum reduced so
as to be broadly isolated from apical mar-
gin; 5th sternum gray pollinose, black; 5th
tergum shiny, black.
Type data.—Musca tenax Linnaeus, lec-
totype @ Linnaean Collection, London, la-
belled ‘“‘tenax 41” and “‘Lectotype, Musca
tenax Linnaeus, Thompson et alia 1981.”
Eristalis columbica Macquart, lectotype
? British Museum (Natural History), Lon-
don, labelled “‘Holotype”’ [red circular type
label], “‘ex. coll. Bigot, Pres. by, G. H. Ver-
rall, B. M. 1894—234;” “‘Eristalis, col-
ombicus, 2 Macq.” [in Macquart’s hand]
and “E. columbicus 2, Columbia Macq.”
[Bigot determination label].
Neotropical material examined.—MEX-
ICO. Aguascalientes: 1 Dec 1909, E C.
Bishopp (1 6 1 2 USNM). Baja California
Norte: Ensenada, 6 Sep 1958, P. H. Arnaud,
jr. (1 2 USNM); Estero Beach, 9 km south
of Ensenada, sea level, 5 Jul 1973, P. H.
Arnaud, Jr., 5 Jul 1973 (1 6 USNM). Chia-
pas: San Cristobal las Casas, 7100 ft., 1—5
Aug 1966, D. E. Breedlove (1 6 USNM);
““Las Casas,” 27 Apr 1945, A. J. Sharp (1
2 USNM). Federal District: 7/8/10, E C.
Bishopp; Federal district: ““7+8 10” (1 @
USNM); Coapa, 25 Aug 1970, E. G. Smyth
dd 2 USNM); Mexico:Gity:; 17° Sep (i 2
USNM); same data, but Crawford (1 @
USNM). México: 20 May 1922, E. G.
Smyth (1 ¢6 1 2 USNM); Atzcaap’co, 31
Aug 1922, E. G. Smyth (3 2 USNM); Teo-
tihuacann, Pyramid to the sun, 27 Dec
1970, P. H. Arnaud (1 6 USNM). Michoa-
can: Morelia, Jun 1965, N. L. H. Krauss (1
2 USNM); Velez Sarssfield, 9 Sep 1926,
M. R. Riesel (1 2 USNM). Nuevo Leon:
Villa de Garcia, 25 May 1975, J. Abercrom-
bie (1 2 USNM). Veracruz: 5 miles south-
west of Perote, 29 Feb 1972, E Parker &
D. Miller (1 6 1 2 USNM); 9 miles south-
west of Toluca, 27 Nov 1965, M. W.
McFadden (5 36 3 2 USNM). GUATE-
MALA. Guatemala: Puerta Parada, 14.5
km east Guatemala City, 16 May 1979 (1
6 USNM). BRAZIL. Paranda: Curitiba, 27
Dec 1936, Westermann (12 USNM); Santa
Catarina, Nova Teutonia, 27°11’S 52°23W,
300-500 m, E Plaumann, 19 Feb 1940 (3 6
USNM), Apr 1947 (1 2 USNM); Nov 1971
(12 6 12 2 USNM), Nov 1964 (2 6 8 92
USNM). Rio Grande do Sul: Pelotas, C. M.
Biezanko, 19 Jan 1956 (2 2? USNM), 29,
30 Sep 1961 (5 6 4 2 USNM), 27, 29 Oct
1960 (2 6 4 2 USNM), 31 Oct 1959 (1 3
3, 2 USNM), 5; 1I-Nov 1956 6G '6-42
USNM), 3, 7 Nov 1960 (8 6 6 2 USNM).
URUGUAY. Montevideo, J. Tremoleras (1
36 1 2 USNM). ARGENTINA. Buenos Ai-
res: Bahia Blanca, 29 Jan 1922, D. S. Bul-
lock (2 ¢6 USNM); Rio Santiago, Palo
Blanco Berisso, 3 Dec 1979, C. M. & O. S.
Flint; Baradero, Rio Parana de las Palmas
Lima, 16 Dec 1979, C. M. & O. S. Flint (1
6 USNM); Balneario Municipal, Baradero,
15 Dec 1979 (1 2 USNM); Buenos Aires,
Jun 1928, A. Copello (4 6 2 2 USNM);
La Plata, 20 Apr 1927, Kisluk (1 36
USNM); Zalaya, Nov 1939, J. B. Daguere
(1 6 2 °2°USNM); Capital, Feb 1939) 378:
Daguerre (1 2 USNM); 7, 24 & 25 Feb
1962, W. L. Jellison (1 6 3 2 USNM);
Azul, 7 -25 Feb 1962, W. L. Jellison (1 3
3 2 USNM); Villa Eliza, 15-29 Dec 1979
(1 6 MZUSP); Villa Eliza, Ao. Carnaval,
3 Dec 1979, C. M. & O. S. Flint (4 6 1 2
USNM); Catamarca: Andalgala, 20 & 28
Oct 1972, J. L. Neff (2 6 2 2 USNM);
Entre Rios: Pronunciamiento, Apr & Dec
1966, E Walz (3 ¢6 3 2 USNM); Neuguen:
Rio Alumine, 9 km north Alumine, 27 Feb
1978, C. M. & O. S. Flint (1 2 USNM);
Ao. Chapelco Chico E. S. M. d. 1. Andes,
25 Feb 1978, C. M. & O. S. Flint (1 6
USNM). CHILE. Southern Chili, 25 Jan
1907, M. J. Rivera (5 ¢ 5 2 USNM). Acon-
cagua: ConCon, 16 Dec 1950, Ross &
Michelbacher (1 ¢ USNM). Antofagasta:
Nov 1932, A. Pirion (1 6 USNM). Arau-
cania: Angol, D. S. Bullock (1 6 3 @
USNM); 11; 23 Mar 1925 @ 6 4.2
USNM), 15 Jun 1933 (1 2 USNM), 22 Nov
VOLUME 99, NUMBER 2
1951 (1 2 USNM). Cautin: Rio Cautin, Ca-
jon, 3 Jan 1966, Flint & Cekalovic (1 2
USNM). Concepcion: Concepcion, Dec
1926, R. & E. Shannon (1 ¢ 1 2 USNM);
Hualpencillo, 31 Dec 1965, Flint & Ceka-
lovic (1 6 USNM). Llanguihue: Puerto Va-
ras, Dec 1926, R. & E. Shannon (1 @
USNM). Maule: Pellines, south of Consti-
tucion, 16 Dec 1976, A. B. Gurney (1 @
USNM). Osorno: Anticura, 1 km west of,
430 m, 11-12 Feb 1978, W. N. Mathis (4
6 1 2 USNM); Aguas Calientes, 1 km
southest of, 530 m, 7-8 Feb 1978, W. N.
Mathis (1 6 USNM); Pucatrihue, 27—30
Jan 1978, W. N. Mathis (1 2 USNM). San-
tiago: Santiago (1 ¢6 USNM); EI Portezue-
lo, 7 km north of Santiago, 500 m, 22—25
Oct 1981, D. & M. Davis (1 2 USNM);
near Pta. Yeso, ca. 70 km southeast Santi-
ago, 1250 m, 27-28 Oct 1981, D. & M. Da-
vis (1 2 USNM); Santiago, “‘10.K.N.09”
29 Jan 1932, Kisliuk & Cooley (1 @
USNM), 29 Feb 1932, Kisliuk & Cooley (1
2 USNM); Valdivia: Rio Bueno, 8 miles
east of, 15 Jan 1951, Ross & Michelbacher
(1 6 USNM). Valparaiso: Valparaiso, A.
Faz (1 6 1 2 USNM), 5 Aug, Cockerell (1
36 USNM); Quillota, Quintolaurel, Kisliuk
& Cooley (1 2 USNM).
Distribution.—Cosmopolitan. Eristalis
tenax is rare in collections from the Neo-
tropics. In temperate areas the species has
spread everywhere humans have. In the
Americas south of the United States, Ervis-
talis tenax is amphitropical, ranging south
to Guatemala and north to Southern Brazil.
Discussion.—Eristalis tenax is a mimic
of the honey bee (Apis mellifera L.). Much
has been written on the species as it has
been used as an experimental subject in var-
ious laboratory studies, a pollinator in
greenhouse, and the basis of the Bugonia
myths.
Subgenus Eoseristalis Kanervo
Eoseristalis Kanervo, 1938: 12. Type spe-
cies, Eristalis cerealis Fabricius (Orig.
des.).
217
Eye without contrasting pile fasciae; aris-
ta usually sparsely pilose basally; meron
bare.
The subgenus Eoseristalis includes some
60 species, mainly distributed in the north
temperate regions (39 Palaearctic species,
20 Nearctic species), with fewer species in
the cooler areas of the Afrotropics (4 spe-
cies), Orient (11 species) and the Neotrop-
ics (5 species).
Eristalis (Eoseristalis) alleni Thompson,
new species
(Fig. 1, habitus; 8, male genitalia)
Male.—Head: Black; face brownish or-
ange except for black oral margin and a
narrow medial black vitta which extends
vertically above tubercle, brownish orange
pilose, dull grayish white pollinose, with
pollinosity thin, not obscuring ground col-
or; gena grayish white pollinose; frontal
and vertical triangle brownish orange pol-
linose, black pilose; eye brown pilose on
dorsal %, yellow on ventral %; occiput sil-
very white pollinose, yellow pilose, with
some black pile on dorsal 4%; antenna or-
ange except darker brown on dorsoapical %
of basoflagellomere; arista orange, distinct-
ly pilose, with pile about % as long as ba-
soflagellomere width; eye contiguity short,
about as long as ocellar triangle.
Thorax: Black, grayish brown pollinose;
scutum with very indistinct pattern of
darker pollinose narrow submedial and
broader interrupted sublateral vittae, exten-
sively black pilose, with orange pilose mar-
ginally (anteriorly, laterally and posterior-
ly); scutellum dark orange, dull, black pi-
lose except apical margin yellow pilose;
pleuron orange pilose except black pile dor-
soposteriorly on anepisternum and posteri-
omedially on katepimeron; metasternum
black pilose. Legs: Orange except black
coxae, trochanters and basally on femora;
orange pilose except black pilose on mid
and hind coxa, dorsoapical % of hind femur,
dorsoposteriorly on hind tibia; anterior spi-
racular fringe light brown; posterior spirac-
ular fringe dark brown; halter brownish or-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 6-10. Male genitalia, lateral view. 6, Eristalis persa. 7, E. croceimaculata. 8, E. alleni. 9, E. bellardii.
10, E. circe.
ange, black head; plumula brown; squama
large, orange brown, with black margin and
fringe; epaulet black and orange pilose;
basicosta black pilose; wing bare, light
brown.
Abdomen: Ist sternum black, gray pol-
linose, yellow pilose; 2nd thru 4th sterna
orange, rarely black medially, yellow pi-
lose; Ist tergum black, gray pollinose, yel-
low pilose; 2nd tergum orange except black
narrowly apicomedially, orange pilose ex-
cept black on apical 4%; 3rd tergum orange
except small black basomedial macula,
black pilose except orange on basal % and
laterally; 4th tergum orange except for very
small black basomedial macula, black pi-
lose except for a few scattered intermixed
orange pile basally and laterally; genitalia
black, dull, yellow and black pilose.
Female.—Similar, but darker; front or-
VOLUME 99, NUMBER 2
angish brown pollinose, black pilose, about
.40 times as wide as head at antenna, ta-
pering to about .20 times as wide dorsally;
fore femur sparsely black pilose posteriorly;
mid and hind femur extensively black pi-
lose; sterna more extensively brownish
black; terga more extensively black pilose,
2nd tergum black pilose on apicomedial %,
3rd tergum entirely black pilose, 4th tergum
extensively black pilose, only orange pilose
laterally; 5th tergum orange basally, black
on apical %, dull pollinose except shiny api-
cally; 5th sternum black, shiny except pol-
linose apically and laterally.
Type data.—Holotype ¢ from COSTA
RICA, Heredia, Braulio Carrillo National
Park, Estacion Barva, 2500 m, L-N 233400
523200, Mar 1990, G. Rivera (INBI-
OCRI000164804), deposited in Instituto
Nacional de Biodiversidad, Santo Domin-
go.
Paratypes: COSTA RICA. Same local-
ity as the holotype but with the following
dates and collectors: Sep 1989, G. Rivera
(1 2 INBIOCRIO00111222); Nov 1989, A.
Fernandez (1 2 INBIOCRI000143385
(USN M)); Dec 1989, A. Fernandez (1 6
1 2 INBIOCRIO00203524 (USNM), INB
IOCRI000290037); Jan 1990, G. Rivera
(43 62 INBIOCRIOO0174715, INBIOCRI
000174710, INBIOCRI000206578
(USNM), INBIOCRIO00206836, INBIOCR
1000192229, INBIOCRI000192230
(USNM), INBIOCRI000206813 (USNM), I
NBIOCRIO000206585, INBIOCRI00020658
0, INBIOCRI000192224); Feb 1990, A
Fernandez (INBIOCRIO00191262-3, INBI
OCRI000191265-6, INBIOCRIO00191268-
73, INBIOCRI000191287); Feb 1990, G.
Rivera (17 36 24 2 INBIOCRI000154527,
INBIOCRI000202034-5, INBIOCRIO0020
2078-9, INBIOCRI000202082, INBIOCRI
000202084-5, INBIOCRI000202087, INBI
OCRIO000202088 (USNM), INBIOCRIO002
02089-90, INBIOCRIO00202097, INBIOC
RI000202105, INBIOCRIO00202108-110, I
NBIOCRIO00202114 (USNM), INBIOCRI
000202115-6, INBIOCRIO00202118, INBI
OCRIO00202120-1, INBIOCRI00020212
219
5-8, INBIOCRI000202130, INBIOCRIOOO
202131 (USNM), INBIOCRI000202132, I
NBIOCRIO000202134-6, INBIOCRI000202
140-1, INBIOCRI000202143-5, INBIOCRI
000202162 (USNM), INBIOCRI00020216
8-9, INBIOCRI0O00202173); Mar 1990, G.
Rivera (2 6 INBIOCRI000164804
(USNM), INBIOCRI000169585); Mar
1990, A. Fernandez (9 2 INBIOCRIO0016
5148, INBIOCRIO00165153, INBIOCRIOOO
164394, INBIOCRI000164327 (USNM), I
NBIOCRIO00164461 (USNM), INBIOCRI
000169474, INBIOCRIO00169479, INBIO
CRI000169427, INBIOCRIO0O00169435).
Heredia: Braulio Carrillo N. P., Transecto,
2050-2600 m, Oct 1989, R. Aguilar & M.
Zumbado (1 ¢ 1 2 INBIOCRIO00131440-
1). Alajuela: Volcan Pods, 9000 ft., 26 Aug
1966, R. D. Akre (1 2 USNM); Volcan
Passy21 Feb 11980; Ae: Waverty1@s G2?
CNC). Cartago: Volcan Irazu, 15 Jul 1965,
G. Fuentas (1 6 USNM). San Jose: Villa
Mills, 25 May 1979, L. Laverty (1 2 CNC),
10 Aug 1979, L. Laverty (1 2 CNC); Ran-
cho Rudondo, 1 Dec 1959, A. Wille (1 ¢
WSU); Cerro de la Muerte, 6 km w Villa
Mills, Inter-Am H’wy, 3340 m, on flowers
of Seneico sp. #234, 24 Feb 1972 0830-
1100 hours, collector E. R. Heithaus
(#10773) (1 6 BMNH), 2-3 Jan 1972, E.
R. Heithaus, on flowers of Seneico oeres-
tedianus (3 ° USNM), 25—26 Jan 1972, E.
R. Heithaus, on flowers of Seneico oeres-
tedianus (1 6 7 2 USNM), 23-25 Feb
1972, E. R. Heithaus, on flowers of Seneico
sp. (6 6 6 2 USNM, 1 ¢ CNC), 21 Jul
1971, E. R. Heithaus, on flowers of Seneico
sp. (1 6 USNM), 25 Jul 1971, E. R. Hei-
thaus, on flowers of Roseaceae (2 6
USNM), 23-26 Oct 1971, E. R. Heithaus,
on flowers of Seneico oerestedianus (4 3 4
2 USNM), 23-29 Nov 1971, E. R. Hei-
thaus, on flowers of Seneico oerestedianus
(17 6 10 2 USNM, 1 6 CNC).
Etymology.—This species is named after
Paul Allen, the co-founder of Microsoft in
recognition of his contributions to the PC
revolution.
Distribution.—Eristalis alleni is known
220
only from the central highlands of Costa
Rica.
Discussion.—Eristalis alleni is similar in
appearance to the males of E. circe and E.
persa, but is readily distinguished by its leg
color.
Eristalis (Eoseristalis) bellardii Jaennicke
(Fig. 9, male genitalia)
Eristalis bellardii Jaennicke, 1867: 400.
Type-loc.: ‘“‘Mexico.”’ Syntypes ¢6 SME
Frankfurt. Williston 1892: 60 (key refer-
ence, description note, Mexico); Curran
1930: 6, 1934: 410 (key references);
Thompson et al. 1976: 101 (catalog ci-
tation).
Eristalomyia rufoscutata Bigot, 1880: 221.
Type-loc.: ““Mexico.”’ Lectotype 6 UMO
(here designated). Synonymy Thompson
ef al 19762 101:
bogotensis of: Giglio-Tos 1893: 4 (descrip-
tion, Mexico); Lynch Arribalzaga 1893:
262 (cit.); Aldrich 1905: 385; Kertész
1910: 214 (in part).
Male.—Head: Black except face some-
times reddish brown laterally; face pale yel-
lowish-white pollinose except for shiny me-
dial vitta which extend % distance to anten-
nal bases, yellow pilose; gena shiny, pale
yellow pilose; frontal lunule reddish brown;
frontal triangle sparsely pale pollinose, yel-
low and black pilose; vertical triangle
sparsely pollinose, black pilose anteriorly,
yellow and black pilose posteriorly; occiput
densely white pollinose on ventral *%4, very
sparsely pollinose on dorsal %, white pilose
becoming more yellowish on dorsal %, with
a few black cilia on dorsal %; antenna
black; arista bare, orange; eye pile brownish
yellow, white ventrally; eye contiguity
long, about 1.5 times as long as vertical tri-
angle.
Thorax: Black, yellow pilose except
black pilose on disc of scutellum and rarely
with a few black hairs on postalar callus,
very sparsely dull pollinose; plumula, halter
and squama pale white to yellowish; spi-
racular fringes brownish yellow; scutellum
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
shiny dark reddish brown. Wing: Epaulet,
tegula yellow pilose; basicosta black pilose;
hyaline, bare except for a few microtrichia
anterobasally on alula. Legs: Extensively
black, pale orange only on apices of fem-
ora, basal % to % of tibiae and on mid bas-
otarsomere, pale yellow pilose except black
pilose on anterior surface of fore and mid
femora and posterior surface of hind femur.
Abdomen: Venter ranging from entirely
brownish black to extensively reddish or-
ange apically, yellow pilose, shiny except
lst sternum sparsely grayish pollinose. Dor-
sum yellow pilose except black pilose api-
comedially on terga; 1st tergum black; 2nd
tergum dull, orange on lateral %, black on
medial %, with apical margin yellow; 3rd
tergum same as 2nd except for indistinct
shiny medial fascia,; 4th tergum orange on
basolateral %, black on apicomedial %, ex-
tensively dull pollinose, with shiny medial
fascia, apical margin yellow; male genitalia
black, shiny.
Female.—Similar; front pale yellowish-
white pollinose, yellow pilose on ventral %4,
black pilose on dorsal %4, about .45 times as
wide as head at antenna, tapering to about
.30 times as wide dorsally; 1st tergum black
medially, orange laterally; 3rd tergum or-
ange only on basolateral % and along apical
margin, elsewhere black; 4th tergum black,
dull pollinose except for broad shiny meidal
fascia; 5th tergum black, shiny medially,
dull pollinose on base and apex.
Type data.—Eristalis rufoscutata Bigot,
lectotype 6 in The Natural History), Lon-
don, labelled with ‘“‘ex. coll. Bigot, Pres. by,
G. H. Verrall, B. M. 1894-234.”’ There are
another 2 males in The Natural History Mu-
seum and 3 more in the Verrall/Collin Col-
lection at Oxford, all of which are labelled
paralectotypes.
Material examined.—MEXICO. no fur-
ther data (1 6 BMNH). Chihuahua: Sierra
Madre, Head of Rio Piedras Verdes, about
7300 ft., C. H. T: Townsend, 21 Jul—11 Sep
(1 6 1 2 USNM), 27 Jun (1 2 BMNH), 3
Jul (2 2 BMNBH), 17 Jul on Rhus glabra (1
2? BMNH); 21 Jul (1 6 2 2 BMNH), 23
VOLUME 99, NUMBER 2
JulegiS73e2 BMNH);'29°Aug @ ¢ 2°29
BMNH), 10 Sep ‘‘flo 273” [=flower
#273?] (1 6 BMNH), 11 Sep on Solidago
trinervata (1 6 BMNH), no date (1 6
BMNH); Durango: El Salto, 10 miles W,
9000 ft., 10 Jun—10 Aug, J. E McAlpine &
J. E. Martin (94 ¢ 105 2 CNC); Buenos
Aires, 10 miles west La Ciudad, 21 Apr
1961 (1 6, 2 2 CNC); Navios, 26 miles
east El Salto, 8000 ft., (3 6, 1 2 CNC); La
Ciudad, 10 miles west along Mexico high-
way #40, 8 Nov 1970, D. E. Breedlove (4
6 USNM); Ciudad, 8100 ft., Forrer (4 6 6
2 BMNH); Huachichilas, 3 miles north,
north base of Las Tabletas, Cerros Huehue-
to, on lumber road, 53 miles north of Coy-
otes, 9000 ft., on flowers Eupatorium and/
or Stevia, 5 Nov 1970, D. E. Breedlove (1
3 USNM). Jalisco: Volcan Tequila, 10-14
km SSW of Tequila, 2134 m, 8 Sep 1974,
Breedlove (1 d CAS). Mexico: Mexico
City, Jun 1918, J. Muller (1 6 USNM);
Morelos: Tres Cumbres, 3 road miles south
of, 9000 ft., at flowers of Lopezia minitata,
16 Jan 1966, D. P. Gregory (1 6 USNM).
Oaxaca: (1 6 BMNH). Chiapas: San Cris-
tobal de las Casas, 11000 ft., 4 Aug 1962,
H. E. Milliron (1 2 CNC); 7200 ft., 25 May
1969, Mason (1 ¢ 1 2 CNC); 7200 ft., 16
May 1969, Mason (1 ¢ CNC); 7087 ft., 16
May 1969, R. V. Peterson (2 6 CNC); 7000
ft., 21 May 1969, H. Teskey (4 6 2 2
CNC); San Cristobal de las Casas, 5 miles
west, 10 May 1969, J. E. Martin (1 ¢o
CNC); San Cristobal, 10 km South, 21 May
ISSINEMeséeO: S. Flint G' ¢ 1 &
USNM); Cerro Huitepec, west San Cristo-
bal de las Casas, 2591 m. 23 May 1972,
Breedlove (1 6 USNM); San Cristobal de
las Casas, 7100 ft., 3 Aug 1966, Breedlove
@°¢ CAS)
Distribution.—Eristalis bellardii is re-
stricted to southwestern United States and
Mexico, where it ranges from Chihuahua to
Chiapas. The northern-most record and
only USA record for E. bellardii is from
southeastern Arizona (Portal). Eristalis bel-
lardii, E. circe and E. persa are sympatric
in Chiapas; E. bellardii and E. circe are
221
broadly sympatric from Durango to Chia-
pas.
Discussion.—Eristalis bellardii repre-
sents the northern component of the E. bo-
gotensis Super species and is distinguished
by pale pile on ventral parts of the thorax
and coxae and the more extensive red col-
oration on the abdomen.
Eristalis (Eoseristalis) bogotensis Macquart
(Fig. 13, male genitalia)
Eristalis bogotensis Macquart, 1842: 112.
Type-loc.: Colombia, Santa Fe de Bogo-
ta. Lectotype 2 MNHN, Paris (here des-
ignated). Williston 1886: 319 (catalog ci-
tation); Wulp 1882a: 79, 1882b: 129 (Ar-
gentina); Lynch Arribalzaga 1892: 253,
1893: 262 (description, Argentina);
Aldrich 1905: 385 (catalog citation); Breé-
thes 1907: 293 (catalog citation); Kertész
1910: 214 (catalog citation); Hull 1925:
308, pl. 2:15 (mesonotum figured, key
reference, description, Bolivia, Peru);
Fluke 1957: 131 (catalog citation, in
part); Gaunitz 1969: 75 (male genitalia
figured); Thompson et al. 1976: 101 (cat-
alog citation)
Eristalis assimilis Macquart, 1846: 257.
Type-loc.: ‘‘Colombia.”” Syntypes 6
Zurich (lost?). Macquart 1850: 499 (in
Collection at Lille, Colombia); Schiner
1868: 361 (descriptive note); Williston
1886: 319 (catalog citation); Kertész
1910: 213 (catalog citation); Hull 1925:
308, pls. 1:7, 2:12 (abdomen figured,
head figured, key reference, description,
Peru); Curran 1930: 6, 1934: 410 (key
reference); Fluke 1957: 130 (catalog ci-
tation, in part); Etcheverry 1963: 34 (syn-
onymy, Chile?). Synonymy Thompson,
et al, 1976: 102:
Male.—Head: Black except face brown-
ish laterally; face shiny except sparsely
white pollinose laterally and ventrad anten-
na, white pilose; gena shiny, white pilose
posteriorly; lunule brownish orange; frontal
triangle sparsely white pollinose laterally,
mainly black pilose, with some white pile
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 11-15.
14, P. eristaloides. 15, E. stipator.
intermixed laterally; antenna black, black
pilose; arista bare, brownish black; eye pile
brown; eye contiguity short, as long as ver-
tical triangle, about % as long as frontal tri-
angle; vertical triangle gray pollinose,
mainly black pilose, with a few yellow
hairs intermixed; occiput densely white pol-
linose, white pilose with some black cilia
on dorsal ¥;.
Thorax: Black except orange humerus,
postalar callus and scutellum; mesonotum
gray pollinose, mainly yellow pilose with
Male genitalia, lateral view. 11, Eristalis gatesi. 12, Palpada semicircula. 13, E. bogotensis.
black pile intermixed; pleuron sparsely gray
pollinose, yellow pilose except black pilose
ventrally; postalar callus yellow pilose; scu-
tellum shiny, yellow and black pilose; squa-
ma yellowish orange; spiracular fringes
brownish yellow; halter yellow. Wing:
Bare, hyaline; tegula yellow and black pi-
lose. Legs: Coxae and trochanters black,
black pilose; femora black except yellow
apex, yellow and black pilose; tibiae yellow
on basal % to %, black apically, yellow pi-
lose; tarsi black, yellow pilose.
VOLUME 99, NUMBER 2
Abdomen: Ist tergum black, black pol-
linose, yellow pilose; 2nd tergum orange
except for medial black T-shaped macula,
dull pollinose, yellow pilose except for a
few apicomedial black hairs, with apical
margin (incisure) yellow; 3rd tergum or-
ange except black medially and apical mar-
gin (incisure) yellow, dull pollinose except
for shiny medial fascia, yellow pilose ex-
cept for black pile apicomedially; 4th ter-
gum black except orange basolaterally and
yellow apical margin (incisure), shiny ex-
cept dull apicomedially, yellow pilose ex-
cept with a few black hairs apicomedially;
sterna black, shiny, yellow pilose; genitalia
black, shiny, yellow pilose.
Female.—Similar; front grayish white
pollinose laterally, more brownish black
pollinose medially, white pilose laterally,
yellow and black pilose medially, about .40
times as wide as head at antenna, tapering
to about .25 times as wide dorsally; abdo-
men with black medial area more extensive,
5th tergum and sternum black, shiny, yel-
low pilose.
Type data.—Eristalis bogotensis Mac-
quart, lectotype 2 #1673 in box 54 of Mac-
quart Collection, Museum National d’ His-
torie Naturelle, Paris, labelled ““No 1144,
Eristalis, bogotensis’” and “‘E. bogotensis
Macq., Bogota.’ There are no original Big-
ot/Macquart labels either in Paris, Oxford
nor London. There are 3 other females in
London and 2d 1 2 in Oxford. All are la-
belled as paralectotypes.
Eristalis assimilis Macquart, types (6 2)
in the collection of Marquis de Bréme. 1 2
with Macquart/Bigot label in BMNH; an-
other listed for Lille.
Material examined.—COLOMBIA. Meta
Dist., 1932, B. Guevara (1 6 1 2 USNM);
Bogota, B. Guevara (50 6 26 2 USNM);
Bogota, Dr. A. Balfour, Feb—Apr 1915 (5 @
BMNH), Dec 1912 (2 d 1 2 BMNH); Bo-
gota, 12 miles southeast, 2930 m, 13 Mar
£955,,8> 1 Schilinger dE: S.,Ross_(h 6
USNM); Volcan Galeras, Narino, 2400 m,
13 Jan 1959, J. FE G. Clarke (1 2 USNM);
Atrat Valley, Boca de Arquia, May—Jun
223
1914 (7 6 4 2 BMNH). ECUADOR. An-
tisamilla to Pinatura, 11000 ft., Ed Whym-
per (2 2 BMNH). Quito, Santa Catalina
Expt. Station, 2780 m, 4 Feb 1971 (3 6 7
2 USNM); Quito, 2850 ft., E Campos R.
(i 6 USNM). Chimborazo: Tixan, 8 mi
northeast of, 14 Feb 1955, E. I. Schlinger
& E. S. Ross (7 d 1 2? CAS, USNM); Lago
Zurucuchu, 11 miles west of Cuenca, 16
Feb 1955, E. I. Schlinger & E. S. Ross (2
36 8 2 CAS, USNM). Tungurahua: Am-
bato, 14 mi northeast of, 2700 m, 8 Feb
1955," E, da Schlinger_ &?E-7S: Ross (i °°
222); Ambato, Nov 1965, J. Foerster (2 3
2 2 CNC). Carchi: Troya, 11-13 Jun 1965,
L. Pena (3 6 CNC); Tulcan, 2800m, 27 Jun
1965, L. Pena (1 d6 2 2 CNC); El Angel,
2700m, 23-25 Jun 1965, L. Pena (7 6 11
2 CNC); Tulcan, 10 km southwest of, 2900
m-, L..Pena @ 6VCNG); Ganar El: dambo;
2800 m, 4—7 Mar 1965, L. Pena (2 6 3 9
CNC). Pichincha: Pomasqui, 2200 m, 6 Jun
1965, L. Pena (1 6 3 2 CNC); Valle de
Machachi, 2900 m, 26 Nov 1940, E Cam-
pos R. (2 6 9 2 USNM); Machachi,
9-10000 ft., Ed Whymper (1 2 BMNH).
Loja: west of Loja, 2500 m, 25 Mar 1965.
(2 6 10 2 CNC). Azuay: Tarqui, 2800 m,
7-8 Mar 1965, L. Pena (2 2 CNC); Cuen-
ca, 28 km south of, 2500—2800m, 15 Mar
1965, L. Pena (1 6 CNC); Cuena, 2200 m,
10-20 Mar 1965, L. Pena (3 6 CNC);
Cuena, 2 km north of, 2200 m., 14 Mar
1965, L. Pena (1 6 1 2 CNC). PERU. Par-
ish (3 ¢ USNM); Tinco?, Aug 1922, Cock-
erell (1 ¢6 USNM); Orova, 7 May 1914, C.
H. T. Townsend (1 6 USNM); Matucana,
C. H. T. Townsend, 30 Jan 1913 (1 6
USNM). Amazonas: Cerros Calla-Calla, 45
km east of Balsas, 3100 m, 20 Jun 1964, P.
C. Hutchison & K. Wright (1 6 USNM).
Arequipa: 2400 m, Aug—Nov 1936, R. M.
Straw (6 6 7 2 USNM); Arequipa, 1926,
Dr. Escomel (1 2 BMNH); Arequipa, 2500
m, 30 Jul-1 Aug 1971 (1 6 1 2 BMNH);
Cuzco, 3200 m, 20—21 Oct 1962, L. Pena
(5 6 4 2 CNC); Cuzco, Las Salineras, 17
Sep 1972.3. Escalente. (ly db, 1). 2 CNC);
Cuzco, Limatambo, 11 Feb 1979, W. E.
224 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Steiner (2 d6 1 2 USNM); Cuzco, Quispi-
camchis, Huambutio, 2900 m, 1 Sep 1988,
A. Freidberg (1 2 USNM); Cuzco, Uru-
bama, 2900 m, 9 Aug 1971, C & M Vardy
(2 6 BMNH); Cuzco, NW of Cuzco, Barrio
Magisterial, C & M Vardy, 3500 m, 7 Aug
1971 (1 6 BMNB), 8 Aug 1971 (1 6
BMNH); 8 kms S of Cuzco, 3500 m, C &
M Vardy, 6 Aug 1971 (1 ¢6 3 2 BMNH);
Moquequa, Yacango, 8 Oct 1965, J. C.
Hitchcock (1 d6 1 2 USNM). Ancashi:
Huaylas, 2800 m., 4 May 1984, P. Hocking
(1 6 1 9 USNM). Junin: 22 Mar 1974, J.
Alata (1 6 USNM); Huancayo, 3300 m, 13
Sep 1984, P. Hocking (1 2 USNM); Tarma,
8 miles west of, 3500 m, 5 Jan 1955, E. I.
Schlinger & E. S. Ross (1 6 2 2 CAS,
USNM). AHuanco: Panao, 3700 m, 27 Jun
1984, P. Hocking (1 2 USNM). Puno: Lago
Titicaca, 11 Mar 1979, W. E. Steiner (2 ¢
1 2 USNM); Roi, 22 Oct 1965, J. C. Hitch-
cock (1 6 1 2 USNM). Mamara, O. Gar-
leppac:, ‘Coll.’ W. Schnusesd911e3 (Ye
BMNH); Mamara, Apurimac, 3500 m, 14
Feb 1910 (1 2 BMNH). BOLIVIA. Cocha-
bamba: Cochabamba, 10 Oct 1966, B. D.
Burks (1 6 USNM), 14-17 Jun 1942 (3 ¢
USNM); Cochabamba, 2600 m, Schoen-
felder, Apr 1950 (1 2 BMNH), 10 Apr
1949 (1 2 BMNH); Lake Titicaca, 26 Apr
1953, J. A. Munro (1 6 USNM); La Paz,
O. Garlepp c., Coll. W. Schnuse 1911-3 (1
36 BMNH); Altiplano, Pillapi, 70 km east
of La Paz, 3780 m, J. L. Chudley, 11 Apr
1964 (1 6 BMNH), 5 May 1964 (1 6
BMNH); ?Yungas del Palmar, 2000 ft.,
Schoenfelder (1 6 BMNH). ARGENTINA.
Tucumadn: Tucuman, 5 Jul 1917 (1 6
USNM); Tucuman, 18 Apr 1913 (1 3
USNM); Tucuman, Tafi del Valle, 2100 m,
2-3 Dec 1979, C & M Varley (2 2
BMNH); Tafi Viejo, La Toma, 10 Oct 1926
(2 6 1 2 USNM). Mendoza: 5 Jan 1927, F
& M Edwards (1 2 BMNH).
Distribution.—BEristalis bogotensis is re-
stricted to South America, where it ranges
from Colombia to northern Argentina. The
southern-most record of E. bogotensis is
from ‘‘Mendoza Prov.,’”? which means that
E. bogotensis and E. croceimaculata are
broadly disjunct in distribution. However,
more collections for central Chile and Ar-
gentina are desired to prove that this dis-
junction is not an artifact due to lack of
collecting. Records of Eristalis tenax,
which does occur in central Chile, strongly
suggest the disjunction is natural.
Discussion.—Eristalis bogotensis of au-
thors is a superspecies, which includes E.
bellardii (USA, Mexico south to El Salva-
dor), E. bogotensis (Colombia to northern
Argentina) and E. croceimaculata (Chile
and southern Argentina). The component
species are readily distinguished by the ex-
tent of pale (red to yellow) maculae on the
abdomen and pale pile on the thorax, with
the amount of pale color being reduced
clinally. The shape of the superior lobe
[=paramere of some] and surstyle of male
genitalia of all species are also distinctive.
Eristalis (Eoseristalis) circe Williston
(Fig. 10, male genitalia)
Eristalis circe Williston, 1891: 59. Type-
loc.: Mexico, Guerrero, Omilteme, 8000
ft. Lectotype ¢ BMNH, London (here
designated). Giglio-Tos 1893: 321 (de-
scription, Mexico); Aldrich 1905: 385
(catalog citation); Kertész 1910: 215 (cat-
alog citation); Hull 1925: 25 (key refer-
ence, description), 1935: 327 (descrip-
tion, Mexico); Curran 1930: 6, 1934: 410
(key reference); Fluke 1957: 132 (catalog
citation); Thompson et al. 1976: 102 (cat-
alog citation).
Eristalis bombusoides Giglio-Tos, 1892: 4.
Type-loc.: Mexico, Oaxaca. Syntypes
3¢ IMZ, Turin. Synonymy by Giglio-
fos 1893-73:
Eristalis aztecus Hull, 1935: 326. Type-
loc.: Mexico, Real del Monte, 9000 ft.
Holotype ¢6 USNM, Washington. Fluke
1957: 131 (catalog citation); Thompson
et al. 1976: 101 (catalog citation) New
synonym.
Male.—Head: Black; face shiny except
sparsely grayish-yellow pollinose ventrad
VOLUME 99, NUMBER 2
antenna, yellow pilose; gena shiny, yellow
pilose; frontal lunule yellow; frontal trian-
gle brownish-yellow pollinose, yellow and
black pilose; eye yellow pilose vertically,
brown pilose dorsally, with yellow and
brown pile medially; eye contiguity short,
about as long as vertical triangle, vertical
triangle brownish-yellow pollinose, yellow
and black pilose; occiput dense grayish-yel-
low pollinose, white pilose ventrally be-
coming orange dorsally except with black
cilia on dorsal %; antenna brown except or-
ange basoventral % of basoflagellomere,
black pilose; arista distinctly pilose on basal
¥, with pile about as long as 2nd antennal
segment width.
Thorax: Black except orange scutellum,
grayish-brown pollinose; mesonotum with
indistinct medial black pollinose vitta; pile
orange; squama brownish black; spiracular
fringes brown; halter yellow with black
knob. Wing: Bare; hyaline except brownish
basally; tegula orange pilose. Legs: Coxae
and trochanters black, orange pilose; fore
and mid femora black except orange on api-
cal ¥,, orange and black pilose; hind femur
black except orange apex, shiny, orange and
black pilose; fore tibia orange on basal %,
black apically, orange pilose; mid tibia or-
ange on basal %, black apically, orange pi-
lose; hind tibia slightly arcuate, black ex-
cept orange on basal %, black pilose; fore
and hind tarsi black, orange pilose; mid tar-
sus orange on basal 2 tarsomeres, apically
black, orange pilose.
Abdomen: Ist tergum black, grayish-yel-
low pollinose, yellow pilose; 2nd tergum
orange except black medial T-shaped mac-
ula, orange pilose; 3rd tergum orange ex-
cept for black quadrate medial maculae, or-
ange pilose; 4th tergum orange, orange pi-
lose; sterna black, yellow pilose; 3rd and
4th sterna with posterior medial edge form-
ing a triangular tooth; genitalia black, or-
ange and black pilose.
Female.—Dissimilar. Head: Similar,
front brownish pollinose, black and orange
pilose on basal %, orange pilose elsewhere,
about .40 times as wide as head at antenna,
225
tapering to about .25 times as wide dorsally.
Thorax: Similar. Wings and legs: Similar.
Abdomen: Entirely black; ist tergum gray
pollinose, yellow pilose; 2nd tergum shiny
except for apical brown pollinose fascia
which is interpreted medially, yellow pilose
on basal *%, brown pilose apically; 3rd and
4th terga shiny except for subapical brown
pollinose fasciae, brown pilose; 5th tergum
shiny, yellow pilose; sterna black, shiny,
white pilose, without apical tooth.
Type data.—Eristalis circe Williston,
lectotype ¢ in The Natural Museum, Lon-
don, labelled “‘Omilteme, Guerrero, 8000
ft., Jul H. H. Smith,” “Biol. Centr. Amer.,
Dipt.-Syrphidae, EF D. Godman, O. Salvin,
1903-51” and “‘‘Eristalis, circe, Williston”’
[a Williston bordered determination label].
There are another 2 males labelled as para-
lectotypes in London, but there are none in
New York.
Eristalis azteca Hull, holotype ¢ the Na-
tional Museum of Natural History, Wash-
ington, labelled ‘“‘Real del Monte, Mex. El
9,000’, “Coll. by H. T. Vanastrand,”’
“Coll. of W. R. Walton,” ‘‘Eristalis, mon-
tanus, Will.,” “‘Eristalis, aztecus, n. sp.”
[determination label in Hull’s hand” and
“Type No., 42076, U.S.N.M.”’ [Red USNM
type label]. The type of aztecus Hull is a
pale immature specimen with brownish or-
ange maculae on 2nd tergum like fenax,
otherwise the specimen agrees well with
my concept of circe.
Material examined.—MEXICO. Cerra-
potosi, 4 Mar 1964, J. E Reinert (2 6
USNM); Real del Monte, 9000 ft., H. T.
Vandstrand (1¢ USNM). Durango: La Ciu-
dad, 24 miles west of, 7000 ft., 2 Jul 1964,
J. E McAlpine (1 ¢ CNC); Buenos Aires,
10 miles west of La Ciudad, 9000 ft., 8
May 1961, H. Howden & J. Martin (1 ¢
CNC); El Salto, 10 miles west of, 9000 ft.,
10 Aug 1964 ( CNC), 12 Jun 1965, J.
McAlpine ( CNC), 30 Jun 1964, J. Martin
( CNC). México: Mexico City, J. Muller (1
6 USNM). Morelos: Huitzilac, 25 Jul 1978,
J. Butze (2 6 1 2 UNAM); Tetela del Vol-
can, 15 Jun 1978, G. Arzate (1 2 UNAM).
226
Oaxaca: Ixtlan de Juarez, 13 miles north-
east of, on Rigidella, 25 Jul 1966, Molseed,
Baptista & Kirchanghi (1 ¢6 1 2? CAS); Si-
erra de Miahuatlan, 2 km southeast of San
Jose del Pacifico, 2438 m, 29 Oct 1974,
Breedlove (3 6 CAS, USNM). Chiapas:
San Cristobal, 8 miles northeast of, 7500 ft.,
9 May 1969, H. Teskey (1 d CNC); 6 May
1969, J. E. Martin (1 2 CNC); Mt. Tzon-
tehuitz, 9500 ft., 27 May 1969, H. Teskey
(2 6 4 2 CNC, USNM). EL SALVADOR.
Monte Cristo, 2418 m., 3 Mar 1978, D. R.
Barger (1 6 USNM), 26 Mar D. R. Barger
(4 6 USNM), 25 Apr 1977, D. R. Barger
(1 6 USNM), 24 May 1977, D. R. Barger
(1 6 USNM), 28 Jun 1977, D. R. Barger
(1 6 USNM), 10 Sep 1977, D. R. Barger
(1 6 USNM); 13 Sep 1977, D. R. Barger
(1 6 1 2 USNM), 15 Sep 1978, D. R. Bar-
ger (1 2 USNM).
Distribution.—Eristalis circe is found in
Mexico (north to Durango) and northern
Middle America (El Salvador), is broadly
sympatric with E. bellardii, and occurs with
E. persa in Chiapas.
Discussion.—Eristalis circe and E. persa
are very similar in appearance, but are
readily separated by their leg color. In both
species, the sexes are strongly dissimilar,
the males having orange colored abdomens
and the females having black colored ab-
domens. Most Eristalis species are mono-
morphic, but E. arbustorum Linnaeus and
E. brousi Williston (two north temperate
species) are also dimorphic. Hence, this
species pair may represent the sister group
to the bogotensis super species.
Eristalis (Eoseristalis) croceimaculata
Jacobs
(Fig. 7, male genitalia)
Eristalis croceimaculata Jacobs, 1900: 107.
Type-loc.: Argentina, Tierra del Fuego,
Isla de los Estados, Golfe Saint-Jean. Ho-
lotype 2 IRSNB, Brussels. Jacobs 1906:
69, pl. 3, fig. 3 (habitus, Argentina); Bré-
thes 1907: 293 (catalog citation); Kertész
1910: 216 (catalog citation); Shannon &
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Aubertin 1933: 122 (?bogotensis); Fluke
1957: 133 (catalog citation).
Palpada croceimaculata: Thompson et al.
1976: 104 (catalog citation).
bogotensis of: Lynch Arribalzaga 1892:
253, 1893: 262 (Argentina); Shannon &
Aubertin 1933: 162 (description note,
Chile, Argentina); Stuardo 1946: 128
(catalog citation, Chile); Fluke 1957: 130
(in part); Etcheverry 1963: 34 (synony-
my, Chile)
Male.—Head: Blackish brown except
face brownish yellow laterally; face shiny
except sparsely white pollinose laterally
and ventrad antenna, yellow pilose except
for a few black hairs dorsolaterally; gena
shiny, white pilose posteriorly; lunule yel-
lowish orange; frontal triangle sparsely
white pollinose laterally, mainly black pi-
lose, with some yellow pile intermixed lat-
erally; antenna black, black pilose; arista
bare, brownish black; eye pile brown; eye
contiguity short, % as long as vertical tri-
angle, about % as long as frontal triangle;
vertical triangle gray pollinose, mainly
black pilose, with a few yellow hairs inter-
mixed; occiput densely white pollinose,
white pilose ventrally, becoming yellow pi-
lose dorsally, with some black cilia on dor-
sal ¥;.
Thorax: Black except orange humerus,
postalar callus and scutellum; mesonotum
gray pollinose with indistinct submedial
black pollinose vittae, mainly yellow pilose
with black pile intermixed; pleuron sparsely
gray pollinose, yellow pilose except black
pilose ventrally; postalar callus yellow pi-
lose; scutellum shiny, yellow and black pi-
lose; squama yellowish orange; spiracular
fringes dirty white; halter yellow. Wing:—
Bare, hyaline; tegula yellow and black pi-
lose. Legs: Coxae and trochanters black,
black pilose; femora black except yellow
apex, yellow and black pilose; tibiae yellow
on basal % to %, black apically, yellow pi-
lose; tarsi black, yellow pilose.
Abdomen: Ist tergum biack, black pol-
linose, yellow pilose; 2nd tergum orange
VOLUME 99, NUMBER 2
except for medial black transverse H-shaped
macula, dull pollinose, yellow pilose except
for a few apicomedial black hairs, with api-
cal margin (incisure) yellow; 3rd tergum
black except orange basolaterally and yel-
low apical margin (incisure) yellow, dull
pollinose except for shiny medial fascia,
yellow pilose except for black pile poste-
riorly; 4th tergum black except yellow api-
cal margin (incisure), shiny except dull ap-
icolaterally, black pilose except yellow pi-
lose basolaterally; sterna brownish black,
shiny, yellow and black pilose; genitalia
black, shiny, yellow and black pilose.
Female.—Similar; front grayish white
pollinose laterally except shiny anteriorly,
white pilose, about .40 times as wide as
head at antenna, tapering to about .25 times
as wide dorsally; abdomen with black me-
dial areas more extensive, 3rd tergum en-
tirely black, 5th tergum and sternum black,
shiny, black pilose.
Type data.—Holotype ¢ in the Institut
Royal des Sciences Naturelles des Bel-
gique, Brussels, labelled ““2,’’ “‘107,’’ “‘Ile
des Etats, Argentine, 8. I. 1897,” “‘Eristalis,
#5, crocei-, maculata J,” ‘‘Eristalis, crocei-
maculata J., det. Jacobs,’ ““TYPE [pink],”
‘cf. Expéd. Antaret. Belg., (Belgica) Zool.
(Ins.), 1906 p. 69 -10,” ‘‘Reg. Mus. Hist.
Nat., Belg. I. G. 10131,” and “‘Eristalis cro-
ceimaculata, J.” The type is double-mount-
ed and in good condition, only the antennae
are missing. The apical tarsomere of the
hind leg is mounted separately and labelled
“A. Collart vid. , 1934:, onote de la patte,
poste,rieure, droite d’Eristalis, croceimacu-
tata J.” and “‘Reg. Mus. Hist. Nat., Belg. I.
G. 10131.”
Material examined.—ARGENTINA.
Chubut: Valle del Lago Blanco, J. Kos-
lowsky, 1904-26 (2 6 BMNH). Rio Negro,
Bariloche, 25-28 Oct 1926, F & M Edwards
(1 2 BMNB), Nov 1926, R. & E. Shannon
(10 6 1 2 USNM), 1 Dec 1926, FE & M.
Edwards (1 6 USNM); Lake Gutierrez,
3-14 Nov 1926, F & M Edwards (3 6
BMNH); Lake Nahuel Huapi, Eastern End,
1 Nov 1926, F & M Edwards (3 6 2 9
227
BMNH). Tierra del Fuego, Rio Grande, Es-
tancia Viamonte, P. W. Reynolds (4 6 1 9
BMNH). FALKLAND ISLANDS. Port
Stanley, Nov 1984-Feb 1985 (1 ¢
BMNH),; East Falklands, Stanley area, Maj.
C. Kirke, Nov 1986 (7 6 2 2 BMNH), 1-
15 Dec 1986 (9 6 3 2 BMNB), 14-31 Dec
1986 (2 6 5 & BMNH), Jan 1987 (3 @
BMNH). CHILE. “‘R. N. El Bolson,”’ 30
Nov 1961, A. Kovacs (1 2 BMNH). Aysen:
Chico, Lag. Buenos Aires, 24—31 Dec
1960, L. Pena (3 2 CNC); Puerto Cisnes,
72°40’W 44°45’, 16-28 Nov 1960, L. Pena
(1 2 CNC); Coihaique, Rio Simpson, 23-—
24 Jan 1961, L. Pena (1 2 CNC), 7-9 Mar
1961, L. Pena (1 2 CNC). Magallanes:
Lag. Amarga, Natales, east of Mt. Payne,
200 m, 14-20 Dec 1960, L. Pena (1 6
CNC); Lena Dura, 4 Dec 1932, E. P. Reed
(1 6 USNM). Tierra del Fuego: Punta Are-
nas, 40 km northeast of, 11 Dec 1960, L.
Pena (1 ¢6 CNC); Puerto Williams, Isla
Navarino, 22—29 Nov 1960, L. Pena (1 2
CNC); Punta Arenas, 9-15 Jan 1966, Flint
& Cekalovic (1 2 USNM), 3 Mar 1959, J.
E. E Clarke (1 6 1 2 USNM). Malleco:
Marimenuco, 1100 m., 10—13 Dec 1959, L.
Pena (1 2 CNC). Llanquihue: Puerto Varas,
Dec 1926, R. & E. Shannon (2 6 USNM);
Casa Panque, F & M Edwards, 4—10 Dec
1926 (1 6 2 2 BMNH).
Distribution.—Eristalis croceimaculata
is restricted to south temperate regions of
South America, being found in Chile and
south Argentina. The northern-most record
for E. croceimaculata is from Marimenuco
in Malleco (Chile). The type-locality on
Staten Island makes it one of the southern-
most recorded flower fly. No other erista-
line flower fly is known from so far south.
However, when a Smithsonian group visit-
ed Staten Island in 1971, no flower flies
were found.
Eristalis (Eoseristalis) gatesi Thompson,
new species
(Fig. 2, habitus; 11, male genitalia)
Male.—Head: Face tawny brown except
broadly black along oral margin and with
228 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
black medial vitta extending over tubercle,
brown pollinose, with pollinosity sparse on
black areas, yellow pilose; gena black,
sparsely gray pollinose, yellow pilose pos-
teriorly; frontal lunule orange; frontal tri-
angle black, gray pollinose, yellow pilose
with a few black hairs intermixed; eye pile
brown; eye contiguity long, as long as ver-
tical triangle, about %4 as long as frontal tri-
angle; antenna orange except narrowly
brownish black on dorsal edge of basofla-
gellomere, black pilose; arista orange,
sparsely pilose on basal %2, with 5-6 long
dorsal hairs and more short vertical ones,
with dorsal hairs about as long as 2nd an-
tennal segment width; occiput densely
white pollinose, yellowish white pilose ven-
trally becoming yellow orange dorsally,
with black cilia on dorsal 4.
Thorax: Black; mesonotum gray polli-
nose, without pattern, yellow pilose except
with black pile intermixed posteriorly; pleu-
ron gray pollinose, black pilose except with
yellow pile intermixed on episternum; post-
alar callus black pilose; scutellum brown
pollinose, black pilose; squama black; hal-
ter gray with black knob; spiracular fringes
black. Wing: Fumose basally, hyaline api-
cally, bare; tegula black pilose. Legs: Black,
black pilose except yellow pilose on fore
tibia and fore and mid tarsi; hind femur nar-
row, very slightly arcuate, without short
black apicoventral spinose hairs.
Abdomen:—1st tergum black, black pol-
linose, black pilose; 2nd tergum yellow ex-
cept black T-shaped basomedial macula and
black basolaterally, black pilose except for
a few yellow hairs basomedially; 3rd ter-
gum yellow except for black T-shaped api-
comedial macula, black pilose; 4th tergum
black, black pollinose except shiny medial
macula, black pilose; genitalia black,
sparsely black pollinose, black pilose; Ist
sternum black, gray pollinose, black pilose
except for a few yellow apicomedial hairs;
2rd and 3rd terga yellow except narrowly
black medially, shiny, yellow pilose; 4th
sternum black, shiny, black pilose except
yellow pilose apicomedially.
Female.—Similar except abdomen more
extensively black; front brown pollinose,
brownish orange pilose, about .40 times as
wide as head at antenna, tapering to about
.20 times as wide dorsally; 2nd tergum
black except narrowly yellow apically,
black pollinose except yellow on apical
margin, entirely black pilose; 3rd tergum
yellow except black laterally and medially,
black pilose; 4th tergum as in male; 5th ter-
gum black, black pollinose basally, shiny
apically, black pilose; 2nd thru 5th sterna
black, shiny, black pilose.
Type data.—Holotype d from Costa
Rica, Heredia, Braulio Carrillo National
Park, Estacion Barva, 2500 m, L-N 233400
523200 G. Rivera (INBIOCRIO00139939).
In Instituto Nacional de Biodiversidad, San-
to Domingo.
Paratypes: COSTA RICA. Same locality
as the holotype but with the following dates
and collectors: Sep 1989, G. Rivera (2d I
NBIOCRIO001 11006, INBIOCRIO0011121
9); Oct 1989, G. Rivera & A. Fernandez (5
2 INBIOCRIO00108636, INBIOCRIO0010
8635 (USNM), INBIOCRIO00108634, INB
IOCRIO00108683 (USNM), INBIOCRIOO0O
108684); Nov 1989, A. Fernandez (1 3 IN
BIOCRIO001433650); Nov 1989, G. Rivera
(2 6 1 2 INBIOCRIO00139937 (USNM),
INBIOCRIO00139932, INBIOCRIOO01401
72 (USNM)); Dec 1989, A. Fernandez (1
2 INBIOCRIO00290043); Jan 1990, G. Ri-
vera (2 6 4 2 INBIOCRIO00192196
(USNM), INBIOCRIO00206816, INBIOCR
1000206810, INBIOCRI000206807, INBIO
CRI000174448, INBIOCRI000192232);
Feb 1990, A. Fernandez (1 6 2 2 INBIO
CRIO00191254-6); Feb 1990, G. Rivera (1
3 4 2 INBIOCRI000202149 (USNM), IN
BIOCRI000202137, INBIOCRI000202107,
INBIOCRIO00202139, INBIOCRIO002021
72, INBIOCRIO000202148); Mar 1990, A.
Fernandez (1 2 INBIOCRI000169656
(USNM); Mar 1990, G. Rivera (1 2 INBI
OCRIO00164807); Apr 1989, M. Zumbado
& A. Fernandez (INBIOCRIO00052536).
Limon: Cerro Chirripo, 2 Aug 1987, A. So-
lis (1 2 INBIO0001007910). Alajuela: Vol-
VOLUME 99, NUMBER 2
can Pass, 21 Feb 1980, T. Laverty (1 ¢
CNC);; Volcan Pods, 9000 ft., 26 Aug
1966, R. D. Akre (2 6 USNM, WSU). San
Jose: Cerro de la Muerte, 2 Mar 1980, T.
Laverty (1 2 CNC); Cerro de la Muerte,
10,000 ft., 24 Aug 1970, R. W. Merritt (1
2 WSU); Cerro de la Muerte, 6 km w Villa
Mills, Inter-Am H’wy, 3340 m, on flowers
of Seneico oerestedianus Benth #234, 23 X
1971 0715-1030 hours, collector E. R. Hei-
thaus (#10773) (1 6 BMNH), 3 Jan 1972,
E. R. Heithaus, on flowers Senecio oerest-
qanus (4%6 3 2° USNM, 16° 1-8 CNG),
25-27 Jan 1972, E. R. Heithaus, on flowers
Senecio oerestdianus (5 3 10 2 USNM),
23 May 1972, E. R. Heithaus, on flowers
Senecio sp. (1 2 USNM), 25 Aug 1971, E.
R. Heithaus, on flowers Rosaceae (3 ¢o
USNM), 23 Aug 1971, E. R. Heithaus, rest-
ing on ground, overcast sky 18°C (1 &
USNM), 18 Aug 1971, E. R. Heithaus, rest-
ing on Vaccinium (1 2 USNM), 23-26 Oct
1971, E. R. Heithaus (1 6 4 2 USNM), 24
Nov 1971, E. R. Heithaus (6 ¢ USNM).
Etymology.—This species is named after
William Gates, III, the co-founder of Mi-
crosoft in recognition of his contributions
to the PC revolution.
Distribution.—Eristalis gatesi is known
only from the central highlands of Costa
Rica.
Discussion.—The black and yellow ab-
dominal color pattern of E. gatesi is unique
among eristaline flower flies.
Eristalis (Eoseristalis) persa Williston
(Fig. 6, male genitalia)
Eristalis persa Williston, 1891: 58. Type-
loc.: Mexico, Guerrero, Sierra de las
Aguas Escondidas, 9000 ft. Holotype 2
BMNH, London. Aldrich 1905: 388 (cat-
alog citation); Kertész 1910: 229 (catalog
citation); Hull 1925: 24 (key reference,
description); Fluke 1957: 140 (catalog ci-
tation); Thompson et al. 1976: 102 (cat-
alog citation).
Male.—Head: Black; face shiny except
sparsely grayish-yellow pollinose ventrad
229
antenna, yellow pilose; gena sparsely gray-
ish-yellow pollinose, yellow pilose; frontal
lunule yellow; frontal triangle brownish-
yellow pollinose, yellow and black pilose;
eye yellow pilose; eye contiguity short,
about as long as vertical triangle, vertical
triangle brownish-yellow pollinose, yellow
and black pilose; occiput dense grayish-yel-
low pollinose, with a brown pollinose mac-
ula on ventral %, yellow pilose except with
black cilia on dorsal %; antenna orange,
black pilose; arista distinctly pilose on basal
¥2, with pile about as long as 2nd antennal
segment width.
Thorax: Black, grayish-brown pollinose;
mesonotum with medial and submedial
black pollinose vittae, with vittae indistinct
anteriorly; pile dark orange, intermixed
black and orange on scutellum and postalar
callus; squama and halter brownish black;
anterior spiracular fringe brown, posterior
spiracular fringe brownish black. Wing:
Bare; hyaline except brownish basally; teg-
ula orange pilose. Legs: coxae and trochan-
ters black, orange pilose; fore and mid fem-
ora black except orange on apical %, orange
pilose; fore femur with dense tuft of black
pile on posterobasal 4%; hind femur black,
shiny, mainly orange pilose, with some
scattered black pile intermixed; fore and
mid tibiae orange, orange pilose; hind tibia
slightly arcuate, black except orange basal-
ly and on apical %, black pilose; tarsi or-
ange, orange pilose.
Abdomen: Ist tergum black, grayish-yel-
low pollinose, yellow pilose; 2nd tergum
orange except black medial T-shaped mac-
ula, orange pilose; 3rd and 4th terga orange
except for black quadrate medial maculae,
orange pilose; sterna brownish orange, or-
ange pilose; genitalia black, orange and
black pilose.
Female.—Dissimilar. Head: Similar ex-
cept more extensively black pilose on front;
front about .35 times as wide as head at
antenna, tapering to about .20 times as wide
dorsally. Thorax: Similar except more ex-
tensively black pilose; pleuron extensively
black pilose except anepisternum extensive-
230
ly yellow pilose; mesonotum with posterior
¥% black pilose except narrowly laterally and
posteriorly orange pilose; scutellum and
postalar callus entirely black pilose. Wings
and legs: Similar except without black pile
tuft on fore femur. Abdomen: Entirely black
and black pilose.
Type data.—Holotype ¢ in The Natural
History Museum, London, labelled ‘‘Sierra
de las Aguas Escondidas, Guerrero, 9500
ft., Jul H. H. Smith,” ““Biol. Centr. Amer.,
Dipt.-Syrphidae, F D. Godman, O. Salvin,
1903-51” and ‘‘Eristalis, persa, Williston”
[a Williston bordered determination label].
Material examined.—MEXICO. Santa
Ana, 9 Jan 1957, P. A. B. (1 2 USNM), 16
Oct 1956, P. A. B. (1 6 USNM). Chiapas:
Municipio Las Margaritas, 48 km northeast
Las Margaritas on road to Campo Alegre,
2134 m, 25 Oct 1976, D. E. & J. A. Breed-
love (1 2 USNM). EL SALVADOR. Mon-
te Cristo, 26 Mar 1978, D. R. Barger (1 6
USNM).
Distribution.—Eyristalis persa is known
only from southern Mexico (Chiapas) and
El] Salvador and is sympatric with E. bel-
lardii and E. circe in Chiapas.
Discussion.—Williston and subsequent
authors did not know the male. The species
is dimorphic like E. circe (q.v.). The male
of E. persa is readily distinguished from all
other Eristalis species by having a tuft of
long black pile on the base of the fore fe-
mur.
Eristalis (Eoseristalis) stipator Osten
Sacken
(Fig. 15, male genitalia)
Eristalis latifrons Loew, 1866: 169 (preocc.
Zetterstedt, 1843). Type-loc.: Mexico,
Matamoros. Lectotype 6 MCZ, Cam-
bridge (here designated). Williston 1892:
60 (key reference, Mexico); Giglio-Tos
1893: 5 (description, Mexico); Aldrich
1905: 386 (catalog citation); Kertész
1910: 224 (catalog citation); Hull 1925:
295 (key reference, description); Curran
1930: 6, 1934: 410 (key reference); Fluke
1957: 137 (catalog citation).
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Eristalis stipator Osten Sacken, 1877: 336.
Type-loc.: USA, Colorado, Manitou
Park. Lectotype ¢ MCZ, Cambridge
(here designated). Thompson et al. 1976:
102 (catalog citation).
Eristalis latifrons var. maculipennis Town-
send, 1897: 93. Type-loc.: USA, New
Mexico, Las ‘Cruces... Lectotypein2
USNM, Washington (here designated).
Townsend 1895: 49 (descriptive note);
Aldrich 1905: 387 (catalog citation); Ker-
tész 1910: 224 (catalog citation).
Male.—Head: Black; face densely white
pollinose except shiny medially on tubercle,
white pilose; gena shiny, white pilose pos-
teriorly; frontal lunule brownish black;
frontal triangle white pollinose, shiny api-
comedially, white pilose; vertical triangle
sparsely brownish pollinose, yellow and
black pilose; eye white pilose; eye conti-
guity long, about % longer than vertical tri-
angle; occiput densely white pollinose,
white pilose ventrally becoming more yel-
low on dorsal 4.
Thorax: Black except orange scutellum,
shiny, orange pilose dorsally, yellow pilose
on pleuron; halter yellow; squama white;
spiracular fringes white. Wing: Bare, hya-
line, tegula yellow pilose. Legs: Coxae and
trochanters black, yellow pilose; femora
brownish black except yellow apical Y,, yel-
low pilose except with some black pile an-
teroventrally on mid and hind femora; fore
tibia orange on basal %, brownish black api-
cally, yellow pilose; mid tibia orange on ba-
sal %, brownish black apically, yellow pi-
lose; hind tibia orange on basal 4%, brownish
black apically, yellow pilose except with
black pile intermixed on apical %; fore tar-
sus brown, yellow pilose; mid tarsus yellow
on basal % of Ist tarsomere and basal % of
2nd tarsomere, brown elsewhere, with pale
areas with yellow pile, with yellow and
black pile elsewhere; hind tarsus black, yel-
low pilose except with black pile inter-
mixed anteriorly.
Abdomen: Ist tergum brown, grayish-
white pollinose, white pilose; 2nd tergum
VOLUME 99, NUMBER 2
brown-black medially in form of X-shaped
macula, yellow elsewhere, shiny except for
brown pollinose apicomedial fascia and
white pollinose apical margin (incisure),
yellow pilose on basal % and laterally, black
pilose on apicomedial %; 3rd tergum
brown, with or without medial transverse
orange macula, shiny except narrowly
white pollinose along anterior margin and
apical margin (incisure), yellow pilose; 4th
tergum shiny except narrowly white polli-
nose along anterior margin and apical mar-
gin (incisure), brownish-black except yel-
lowish-white apical margin, white pilose;
genitalia brownish black, shiny, yellow pi-
lose; Ist sternum brown, gray pollinose,
yellow pilose; 2nd sternum brownish me-
dially, yellow laterally, shiny, yellow pi-
lose; 3rd and 4th sterna brownish black,
shiny, yellow pilose.
Female.—Similar; front entirely white
pilose, about .45 times as wide as head at
antenna, tapering to about .30 times as wide
dorsally; 5th tergum and sternum black,
yellow pilose.
Type data.—Eristalis latifrons Loew,
lectotype 6 in Museum of Comparative Zo-
ology, Cambridge, labelled ‘‘Mat,’’ ‘““Loew
Coll,” “Type, 4073” and “‘Eristalis, latif-
rons, m.”’ [in Loew’s hand].
Eristalis stipator Osten Sacken, lectotype
6 in Museum of Comparative Zoology,
Cambridge, labelled ‘‘Manitou, Park,”’
“Osten, Sacken, Coll.,” ‘““Type, 7,885” and
“‘latifrons.”’ There are another 8 specimens
from various localities that have been la-
belled as paralectotypes.
Eristalis latifrons var. maculipennis
Townsend, lectotype 2? in National Muse-
um of Natural History, Washington, la-
belled “‘Las Cruces, 6.7 N.M.”’ and ‘Coll.
Townsend.” No other syntypes were found.
George Byers carefully checked the Snow
Museum collections, where additional ma-
terial should have been deposited. While
this variety was formally named in 1897,
Townsend referred back to his description
of it in his 1895 paper. Hence, the type se-
ries consists of the female mentioned in the
231
1897 as well as those mentioned in the ear-
lier paper. The Las Cruces material of his
1895 paper is undoubtedly what he refers
to as “numerous specimens from Mesilla
valley of the Rio Grande”’ in his 1897 pa-
per.
Material examined.—MEXICO. Chihua-
hua: Sierra Madre, Head of Rio Piedras
Verdes) about), 72300.ft..03), Juls-C. He “T.
Townsend (1d USNM); same locality, but
“9.6” (1 2 USNM). Also, numerous spec-
imens from the United States were exam-
ined.
Distribution.—Eristalis stipator is found
in southern Canada (British Columbia to
Nova Scotia), throughout the United States
and in northern Mexico. The above record
is the southern-most for the species.
OTHER SPECIES
A few other species of Eristalis have
been erroneously reported from the neo-
tropics. Eristalis arbustorum and E. trans-
versa were incorrectly recorded from Ja-
maica (Thompson 1981: 147). Macquart
(1842: 32) described Eristalis quadelupen-
sis from Guadeloupe which was based on
an apparently mislabeled specimen of E.
pertinax (Scopoli)(Thompson 1981: 146).
Likewise, Bigot (1880: 217) described E.
inca from Peru. Verrall (1901: 514; Kertész
1910: 230) noted that the name was based
on a specimen of E. pertinax, an identifi-
cation which has been confirmed recently
(Nielsen, in litt.). The following two Pal-
pada species are treated here as they may
be confused with Ervistalis species due to
their bare katepimera.
Palpada eristaloides Thompson,
new species
(Fig. 14, male genitalia)
Male.—Head: Face brownish yellow,
yellow pollinose except shiny medial vitta,
yellow pilose; gena yellow brown, shiny,
yellow pilose posteriorly; occiput white
pollinose and pilose on ventral %4, more
grayish pollinose on dorsal %4, black pilose
dorsally; frontal lunule yellow; front dark,
232
brownish black pollinose except more yel-
lowish pollinose laterally along eye margin,
black pilose; frontal triangle yellow, yel-
lowish-white pollinose, black pilose; vertex
black, brownish-black pollinose, black pi-
lose; eye black pilose on dorsal %, yellow
pilose ventrally; eye contiguity short, about
¥% as long as vertical triangle; eyes dichop-
tic, separated by distance equal to aristal
width; antenna orange, orange pilose except
for a few long black bristle-like hairs on
2nd segment; basoflagellomere with a large
basoventral sensory pit on mesal surface;
arista brownish black, bare.
Thorax: Black; mesonotum generally
dull black pollinose, with pale grayish-
white pollinose pattern, black pilose medi-
ally, dark brownish-orange pilose anteriorly
and laterally; mesonotal pale areas anteri-
orly and laterally, on transverse suture, and
in form of broad submedial vittae which ex-
tend % distance to scutellum; postalar callus
dark brownish-black pilose; scutellum yel-
lowish orange, slightly brownish black ba-
somedially, black pilose; pleuron gray pol-
linose, brownish-orange pilose; ampulla or-
ange; plumula and halter yellow; squama
brownish black on dorsal lobe and brown
on margin of ventral lobe, yellowish orange
elsewhere; spiracular fringes white. Wing:
Hyaline, bare except microtrichose posteri-
or to stem vein and basoanterior corner of
alula. Legs: Coxae black, gray pollinose,
yellow pilose; trochanters black, shiny ex-
cept fore trochanter gray pollinose, yellow
pilose; hind trochanter with dense apical
tuft of black setulae; fore and mid femora
blackish brown except yellow apically, yel-
low on apical ¥% anteriorly, apical % poste-
riorly, yellow pilose; hind femur arcuate,
blackish brown except yellow on apical %,
yellow pilose with short black spinose hairs
ventrally; tibiae yellow, yellow pilose; hind
tibia with ventromedial carina on basal %,
with apicolateral spinose setal patch; fore
tarsus black except yellow basotarsomere,
yellow pilose; mid and hind tarsi black ex-
cept yellow basotarsomere and basal % of
2nd tarsomere, yellow pilose.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Abdomen: Ist tergum yellow except
brown on medial %, gray pollinose, yellow
pilose; 2nd tergum dull yellowish orange
except black medial T-shaped maculae and
apicolateral corners, yellowish orange pi-
lose except for a few black hairs apicome-
dially; 3rd tergum dull black except large
orange basolateral maculae on basal *% and
yellow apical margin, black pilose except
for a few longer lateral yellow hairs; 4th
tergum dull black except yellow apical mar-
gin, black pilose; 5th tergum black, shiny
on apical %, dull black pollinose on basal
¥%, black pilose; 1st sternum reduced, yel-
low, sparsely white pollinose, yellow pi-
lose; 2nd thru 4th sterna yellow, slightly
more brownish medially and laterally,
shiny, yellow pilose; 5th tergum_ black,
shiny, yellow pilose.
Female.—Similar; front entirely black pi-
lose, dark brown pollinose medially and en-
tirely on dorsal %, light tan pollinose lat-
erally on ventral %, brown pollinose else-
where, about .35 times as wide as head at
antenna, tapering to about .15 times as wide
dorsally. Legs: Darker, basal % of front and
middle femora brown, basal % of hind fe-
mur brown, hind tibia and trochanter with-
out black setal patches or tufts; 5th tergum
brownish black, black pollinose on basal %,
shiny apically.
Type data.—Holotype 6 in Canadian
National Collection, Ottawa, labelled EC-
UADOR: Carchi, 10 km SW Tulcan, 2900
m, 28 Jun 1965 (L. Pefia).
Paratypes: ECUADOR. Carchi, Troya,
2950 m, 11-13 Jun 1965 (L. Pena) 1 6
(USNM); Azuay, Cerro Tinajillas, 3100 m,
18-21 Mar 1965 (L. Pefia) 1 2 (CNC);
Napo, 0°22’S 78°8’W, 3500 m, 4—7 Mar
1976 (G. & M. Wood) 2 2 (CNC, USNM).
Etymology.—This species is named er-
istaloides as it is phenotypically like Eris-
talis in having the katepimeron bare.
Distribution.—Palpada eristaloides is
known only from high elevations in Ecua-
dor.
Discussion.—Palpada eristaloides is dis-
VOLUME 99, NUMBER 2
tinctive with its mesonotal pollinose pattern
and bare katepimeron.
Palpada semicircula Walker
(Fig. 12, male genitalia)
Eristalis semicirculus Walker, 1852: 249.
Type-loc.: Honduras. Lectotype ¢d
BMNH, London (here designated). Wil-
liston 1892: 78 (citation); Wulp 1896:
114 (East Indies [error]), 1899: 52 (cor-
rection of previous error); Aldrich 1905:
389 (catalog citation); Kertész 1910: 235
(catalog citation); Fluke 1957: 143 (cat-
alog citation).
Palpada_ semicirculus: Thompson et al.
1976: 109 (catalog citation).
Eristalis tenuifrons Curran, 1930: 12. Type-
loc.: Panama, Canal Zone, Fort Ran-
dolph. Holotype 2 AMNH, New York.
Curran 1934: 409 (key reference); Fluke
1957: 145 (catalog citation); Thompson
et al. 1976: 102 (catalog citation). New
synonym.
Male.—Head: Black; face white polli-
nose except for shiny narrow medial vitta,
white pilose; gena shiny; frontal triangle
shiny apicomedially, white pollinose else-
where, black pilose medially, white pilose
laterally; vertical triangle gray pollinose an-
teriorly, black pollinose on ocellar triangle,
black pilose; occiput densely white polli-
nose and pilose on ventral %, more brown-
ish black pollinose and black pilose dorsal-
ly; antenna black pilose; scape and pedicel
brownish orange; basoflagellomere orange-
brown on basoventral %, brownish black
elsewhere; arista bare; eye pile short, white;
eyes narrowly dichoptic, separated by about
basal width of arista.
Thorax: Black, extensively yellowish
white pilose, black pilose on scutellum and
postalar callus, with some intermixed black
pile on posterior % of scutum; mesonotum
dull black pollinose except broadly gray
pollinose anteriorly, laterally, across trans-
verse suture and anterior to scutellum; gray
pollinose areas appear as 2 broad anterior
fasciae connected to a large U-shaped mac-
233
ula posteriorly; pleuron gray pollinose, with
pollinosity densest on katepisternum; scu-
tellum yellow except black basal %; halter
orange; plumula white; squama white with
black margin and yellow apical fringe; spi-
racular fringes dirty white. Legs: Black ex-
cept brownish orange femoral-tibial joints,
pale pilose except black pile on anterior and
ventral surfaces of mid tibia, black pilose
on anterodorsal surface of fore and mid
femora, black pilose on apex and apico-
ventral margin of hind femur and black pi-
lose on tarsi; hind femur swollen; hind tibia
with ventromedial carina on apical 2 in ad-
dition to basoventral carina; hind trochanter
with only normally white pile; epaulet
black pilose except for some orange pile;
tegula brown pilose; basicosta black pilose;
wing hyaline, bare except for a few widely
scattered microtrichia apically and posteri-
orly.
Abdomen: Sterna 1, 3 and 4 black, gray
pollinose, white pilose; sternum 2 yellow,
shiny, white pilose; Ist tergum black, gray
pollinose, white pilose; 2nd tergum exten-
sively black, dull black pollinose, white pi-
lose, with large subtriangular shiny yellow
basolateral macula and yellow apical mar-
gin (incisure), with macula narrowly sepa-
rated from lateral margin and occupying ba-
solateral *%4 of tergum; 3rd tergum similar to
2nd, with macula smaller, occupying only
basolateral %, more extensive black pilose
on apicomedial *%4; 4th tergum black except
yellow apical margin (incisure), shiny on
basal *%, dull pollinose apically, black pilose
on apical %, elsewhere black pilose medi-
ally, yellow pile laterally.
Female.—Similar to male (see Curran’s
description).
Type data.—Eristalis semicirculus Walk-
er, lectotype of unknown sex, in The Nat-
ural History Museum, London, labelled
“Type” [green circular type label], ““Ho-
lotype”’ [red circular type label], “‘Hond.,
Dys.,”’ ‘““S. America, Hondura,”’ “‘Eristalis,
semicirculus, Wlk.’’ [in Austen’s hand] and
‘*semicirculus”’ [in Walker’s hand]. A mere
fragment remains of the type of E. semicir-
234
cula Walker, only the thorax and basal two
segments of the abdomen are extant. How-
ever, enough remains to be certain of its
identity.
Eristalis tenuifrons Curran, holotype °
in American Museum of Natural History,
New York, labelled “‘Panama, Canal Zone,
Fort Randolph, Feb 6, 1929, C. H. Curran,”
holotype Eristalis tenuifrons Curran [red
type label in Curran’s hand].
Material examined.—BELIZE. Stann
Creek, Sittes Point, Malaise trap at Possum
Point Biological Station, 22—30 April 1987,
Spangler & Faitoute (57 6 USNM). PAN-
AMA. Darien: Garachine, Feb 1953, ES.
Blanton (1 6 USNM). Canal Zone: Ca-
maron, Fort Kobbe, 22 Jul 1952, FE S. Blan-
ton (1 6 USNM).
Distribution.—Palpada semicircula is
currently known only from Belize and Pan-
ama, but probably will be found throughout
the lowlands of Central America.
Discussion.—Palpada semicircula is a
small eristaline fly with the typical Palpada
appearance of gray pollinose fasciae on the
mesonotum and pale maculae on the ab-
domen. However, the species is immediate-
ly separated from almost all other Palpada
species by the bare katepimeron. Palpada
eristaloides is the only other Palpada spe-
cies with a bare katepimeron and is distin-
guished by its larger size and lack of gray
fasciae on the mesonotum.
ACKNOWLEDGMENTS
I thank Paul Arnaud, Jr., Department of
Entomology, California Academy of Sci-
ences, San Francisco (CAS); Javier Butze,
formerly with Instituto de Biologia, Univ-
ersidad Nacional Autonoma México, Mex-
ico City (UNAM); Brian Pitkin and Adrian
Pont, The Natural History Museum (for-
merly the British Museum (Natural Histo-
ry)), London (BMNH); David A. Grimaldi,
the American Museum of Natural History,
New York (AMNH); Loic Matile, Ento-
mologie, Muséum National d’ Histoire Na-
turelle, Paris (MNHN); Manuel Zumbado,
Instituto Nacional de Biodiversidad, Here-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
dia (INBio); J. R. Vockeroth, Canadian Na-
tional Collection, Agriculture Canada, Ot-
tawa (CNC); Paul Hanson, University of
Costa Rica, San José (UCR); George Byers,
Snow Entomological Museum, Department
of Entomology, University of Kansas, Law-
rence (UKaL); Patrick Grootaert, Institut
Royal des Sciences Naturelles des Bel-
gique, Brussels (IRSNB); and Richard
Zack, Department of Entomology, Wash-
ington State University, Pullman (WSU) for
permission to study material in their care. I
also thank Drs. Neal Evenhuis, Bishop Mu-
seum, Honolulu; J. R. Vockeroth, Canadian
National Collection, Agriculture Canada,
Ottawa; John Oswald, Department of En-
tomology, Smithsonian Institution, Wash-
ington, D.C. (USNM); and Allen Norrbom
and Manya B. Stoetzel, Systematic Ento-
mology Laboratory, USDA, Washington,
D.C., and Beltsville, Maryland, for their
critical reviews of the manuscript.
Appreciation is due also to Steven Falk
(Figs. 1-2), Arthur Cushman (Fig. 4), Can-
dy Feller (Fig. 5) and Lisa Roberts (Figs.
6-15). All figures except the color ones
were prepared at government expense and
are without copyright. The color plate
(Figs. 1-2) was prepared at my expense and
as copyright holder I hereby make these fig-
ures available for non-commerical and sci-
entific use only. Last but not least, acknowl-
edgment is due to the Samuel Wendell Wil-
liston Diptera Research Fund of the Smith-
sonian Institution and the Instituto Nacional
de Biodiversidad for supporting my field
work in Costa Rica.
LITERATURE CITED
Aldrich, J. M. 1905. A catalogue of North American
Diptera. Smithsonian Miscellaneous Collections
46(2), 680 pp.
Bigot, J. M. EF 1880. Diptéres nouveaux ou peu con-
nus. 14e partie, X XI: Syrphidi (mihi), Genre Er-
istalis (Fabr.). Annales de la Société Entomolo-
gique de France (5) 10: 213-230.
Bréthes, J. 1907. Catalogo de los Dipteros de las Re-
pliblicas del Plata. Anales Museo Nacional de
Buenos Aires (3)9: 277-305. (28 Oct.)
Coquillett, D. W. 1910. The type-species of the North
VOLUME 99, NUMBER 2
American genera of Diptera. Proceedings of the
United States National Museum 37: 499-647.
Culotta, E. 1992. Museums cut research in hard times.
Science 256: 1268-1271.
Curran, C. H. 1930. New species of Eristalinae with
notes (Syrphidae, Diptera). American Museum
Novitates 411, 27 pp. (21 Mar.)
Curran, C. H. 1934. The Diptera of Kartabo, Bartica
District, British Guiana. Bulletin of the American
Museum of Natural history 66: 287—532. (30 Jul.)
Curtis, J. 1832. British Entomology: Being illustra-
tions and descriptions of the genera of insects
found in Great Britain and Ireland. Vol. 9, pls.
384—433. London.
Enderlein, G. 1940. Die Dipterenfauna der Juan-Fer-
nandez-Inslen und der Oster-Inseln. Pp. 643-680.
In Skottsberg (ed.), Natural History of Juan Fer-
nandez and Easter Island. Zoology 3.
Erwin, T. L. 1992. A current vision of insect diversity,
pp. 91-97. In Sarukhan, J. and Dirzo, R., comp.,
México ante los de la biodiversidad. 343 pp. Com-
isidn Nacional para el conocimiento y uso de la
Biodiversidad, Mexico
Etcheverry, M. 1963. Descripciones originales, sinon-
imia y distribucion geografica de las especies de
la familia Syrphidae (Diptera) en Chile. Publica-
ciones del Centro de Estudios Entomologicos,
Universidad de Chile 5, 144 pp.
Etcheverry, M. and R. D. Shenefelt 1962. A prelim-
inary study of the genitalia of Chilean Syrphidae.
XI. Internationaler Kongress fiir Entomologie,
Wien 17. bis 25. August 1960. Verhandlungen 1:
207-214.
Fabricius, J. C. 1805. Systema antliatorum secundum
ordines, genera, species adiectasynonymis, locis,
observationibus, descriptionibus. 373 pp., + 30
pp. Reichard, Brunsvigae [=Brunswick].
Fluke, C. L. 1957. Catalogue of the family Syrphidae
in the Neotropical Region. Revista Brasileira de
Entomogia 7: 1-181.
Gamez, R. and I. D. Gauld 1993. Costa Rica: An
innovative approach to the study of tropical bio-
diversity, pp. 329-336. In LaSalle, J. and Gauld,
I. D., eds., Hymenoptera and Biodiversity. xx +
999 pp., C.A.B. International, Redwood Brooks.
Gaston, K. J. and R. M. May 1992. Taxonomy of
taxonomists. Nature 356: 281-282.
Gaunitz, S. 1969. Studien iiber die unterfamilie Eris-
talinae. Der Bau des mannlichen genitalapparants
(Dipt., Syrphidae). Entomologisk Tidskrift 90:
73-99. (15 Aug.)
Giglio-Tos, E. 1892. Diagnosi di nuove specie di Dit-
teri. VI. Sirfidi del Messico (1). Bollettino dei Mu-
sei di Zoologia ed Anatomia Comparata della R.
Universita di Torino 7 (123), 7 pp.
Giglio-Tos, E. 1893. Ditteri del Messico. Pt. 2, Syr-
phidae—Conopidae—Pipunculidae. 80 pp. Tori-
i)
eS)
Nn
no. Also published in Mem. Reale Accademia del-
le Scienze Torino (2) 43: 321-398, 1 pl.
Goffe, E. R. 1946. The genera in Syrphidae of Lioy,
1864 (Diptera). Journal of the Society for British
Entomology 3: 27-30. (25 Jun.)
Gutiérrez, R. 1939. Insectos capturados en Rio Blan-
co. Revista Chilena de Historia Natural 43: 32-
315).
Holden, C. 1989. Entomologists wane as insects wax.
Science 246: 754-756.
Hovore, E T. 1991. INBio: By biologists for biolo-
gists. American Entomologist 37: 157-158.
Hull, EF M. 1925. A review of the genus Eristalis.
Ohio Journal of Science 15: 11—43 [Part 1], 285—
310 [Part 2].
Hull, E M. 1935. Some undescribed species of Eris-
talis from North America in the United States Na-
tional Museum. Journal of the Washington Acad-
emy of Sciences 25: 326-331.
International Commission on Zoological Nomencla-
ture (I.C.Z.N.) 1963. Opinion 678. The suppres-
sion under the Plenary Powers of the pamphlet
published by Meigen, 1800. Bulletin of Zoologi-
cal Nomenclature 20: 339-342.
International Commission on Zoological Nomencla-
ture (I.C.Z.N.) 1993. Opinion 1747. Eristalis La-
treille, 1804, Helophilus Fabricius, 1805, Xylota
Meigen, 1822, and Eumerus Meigen, 1822 (Insec-
ta, Diptera) conserved. Bulletin of Zoological No-
menclature 50: 256—258.
Jacobs, J. Ch. 1900. Diagnoses d’Insectes recueillis
par l’expedition Antarctique Belge. Annales de la
Société Entomologique de Belgique 44: 107.
Jacobs, J. Ch. 1906. Bibionidae, Chironomidae, Cu-
licidae, Tipulidae, Syrphidae, Muscidae, Rhyphi-
dae, Anthomyidae, pp. 67-71, pl. 3, figs. 3-5. In
Belgium, Commission de la Belgica. Expédition
Antarctique Belge. Résultats du Voyage du S. Y.
Belgica en 1897-1898-1899 sous le commande-
ment de A. de Gerlache de Gomery. Rapports
scientifiques. Zoologie, Insectes (Diptéres).
Jaennicke, EK 1867. Neue exotische Dipteren. Abhan-
dlungen herausgegeben von der Senckenbersichen
Naturforschenden Gesellschaft 6: 311—408, pls.
43—44. Also, published separately.
James, M. T. 1947. The flies that cause myiasis in
man. U. S. Department of Agriculture, Miscella-
neous Publication 631, 175 pp.
Janzen, D. H. 1991. How to save tropical biodiversity.
American Entomologist 37: 159-171.
Kanervo, E. 1938. Zur systematik und Phylogenie der
westpalaarktischen Eristalis-arten (Dipt. Syrphi-
dae) mit einer Revision derjenigen Finnlands. An-
nales Universitatis Turkuensis (A) 6(4), 54 pp.
Kertész, K. 1910. Catalogus dipterorum hucusque
descriptorum. Vol. 7, 470 pp. Budapest.
Latreille, P A. 1804. Tableau méthodique des Insec-
tes, pp. 129-200. In Societé de Naturalistes et
d’ Agriculturs, Nouveau dictionnaire d’ histoire na-
turelle, appliqué aux arts, principalement 4 1’ec-
onomie rurale et domestique. Vol. 24: Tableaux
méthodiques d’histoire naturelle. 238 pp. Paris.
Latreille, P. A. 1810. Considérations générales sur
l’ordre naturel des animaux composant les classes
des Crustacés, des Arachnides, et des Insectes;
avec un tableau méthodique de leurs genres, dis-
posés en familles. 444 pp. Schoell, Paris. (23
May)
Linnaeus, C. 1758. Systema naturae per regna tria na-
turae, secundum classes, ordines, genera, species,
cum caracteribus, differentiis, synonymis, locis.
Ed. 10, Vol. 1. 824 pp. L. Salvii, Holmiae
[=Stockholm]. (1 Jan.)
Lioy, P. 1864. I ditteri distributi secondo un nuovo
metodo di classificazione naturale. Atti dell’ I. R.
Istituto Veneto di Scienze, Lettere ed Arti (3) 9:
738-760.
Loew, H. 1866. Diptera Americae septentrionalis in-
digena. Centuria sexta. Berliner Entomologische
Zeitschrift (1865) 9: 127-186.
Lynch Arribalzaga, E 1892. Dipterologia Argentina
(Syrphidae). [part] Anales de la Sociedad Cienti-
fica Argentina 33: 236—254. (Jun.)
Lynch Arribalzaga, EF 1893. Dipterologia Argentina
(Syrphidae). [part] Anales de la Sociedad Cienti-
fica Argentina 34: 241-280. (Jan.)
Macquart, J. 1842. Dipteres exotiques nouveaux ou
peu connus. Mémoires de la Société des Sciences,
de I’ Agriculture et des Arts de Lille 1841 (1): 65—
200, 22 pls. Also published separately as his
“Diptéres exotiques nouveaux ou peu connus,”
vol. 2, Pt. 2, pp. 5-140, 22 pls. Paris.
Macquart, J. 1846. Diptéres exotiques nouveaux ou
peu connus. Supplément. Mémoires de la Société
des Sciences, de |’ Agriculture et des Arts de Lille
(1845) 1844: 133-364, 20 pls. Also published
separately as his ““Dipteres exotiques nouveaux ou
peu connus. Supplément,” pp. 5—238, 20 pls. Par-
is.
Macquart, J. 1850. Animaux invertébrés. Catalogue
du Musee d’ Histoire naturelle de la Ville de Lille.
Tome 2, iv+ 634 pp. Lille.
Macquart, J. 1855. Diptéres exotiques nouveaux ou
peu connus. 5.° supplément. Mémoires de la So-
ciété des Sciences, de |’Agriculte et des Arts de
Lille (2) 1: 25-156, 7 pls. (before 18 Aug.)
May, R. M. 1990. How many species? Philosophical
Transaction of the Royal Society, London 330 (B):
293-304.
May, R. M. 1992. How many species inhabit the
earth? Scientific American, Oct, pp. 42—48.
Meigen, J. W. 1800. Nouvelle classification des
mouches a deux ailes (Diptera L.) d’apres un plan
tout nouveau. 40 pp. Paris. A Facsimile appears
in Hemming, 1945: 121—160. This publication has
been suppressed (I.C.Z.N. 1963).
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Meigen, J. W. 1803. Versuch einer neuen Gattung-
seintheilung der europaischen zweiflugeligen In-
sekten. Magazin fiir Insektenkunde 2: 259-281.
Meigen, J. W. 1804. Klassifikazion und Beschreibung
der europaischen zweiflugeligen Insecten (Diptera
Linn.). Erster Band. Abt. I. pp. i-xxviii + 1-152,
pls. 1-8; Abt. II, pp. i-vi + 153-314, pls. 9-15.
Braunschweig.
Meigen, J. W. 1822. Systematische Beschreibung der
bekannten europaischen zweiflugeligen Insekten.
Vol. 3, x + 416 pp., pls. 22-32. Hamm.
Mik, J. 1897. Einige Bemerkungen zur Dipteren-Fam-
ilie der Syrphiden [concl.] Wiener Entomologis-
che Zeitung 16: 113-119. (31 Mar.)
National Science Board (NBS). 1989. Loss of biolog-
ical diversity: A global crisis requiring interna-
tional solutions. Report NSB-89-171, vii + 19 pp.
National Science Foundation, Washington.
Novacek, M. J. 1992. The meaning of systematics and
the biodiversity crisis, pp. 101-108. Jn Eldredge,
N., ed., Systematics, ecology, and the biodiversity
crisis. xiv + 220 pp., Columbia University Press,
New York.
Osten Sacken, C. R. 1877. Western Diptera: Descrip-
tions of new genera and species of Diptera from
the region west of the Mississippi and especially
from California. Bulletin of the United States
Geological and Geographical Survey of the Ter-
ritories 3: 189-354.
Porter, C. E. 1921. Sobre algunos Sirfidos de Chile.
Revista Chilena de Historia Natural 25: 446-447.
Porter, C. E. 1924a. Algunos Insectos colectados en
el sur del pais por el enor Faminio Ruiz P. Revista
Chilena de Historia Natural 28: 81-82.
Porter, C. E. 1924b. Enumeracion de sirfidos colec-
tados en San Jose de Maipo en Octubre y Novbre.
de 1923. Revista Chilena de Historia Natural 28:
98.
Porter, C. E. 1927. Algunos insectos de Mancera. Re-
vista Chilena de Historia Natural 31: 121-122.
Porter, C. E. 1928. Entomologia Chilena. Primera lista
de insectos de Panimavida. Revista Chilena de
Historia Natural 32: 221-225.
Porter, C. E. 1932. Acerca de algunos insectos Chi-
lenos. Revista Chilena de Historia Natural 36:
190-193.
Porter, C. E. 1934. Insectos colectados en Puente Alto
(1). Revista Chilena de Historia Natural 38: 169—
1 7/ile
Porter, C. E. 1938. Algunos insectos de las provincias
de Atacama y Coquimbo. Revista Chilena de His-
toria Natural 42: 154-155.
Raven, P. H. and Wilson, E. O. 1992. A fifty-year plan
for biodiversity surveys. Science 258: 1099-1100.
Roberts, L. 1988. Hard choices ahead on biodiversity.
Science 241: 1759-1761.
Rondani, C. 1845. Ordinamento sistematico dei generi
italiani degli insetti ditteri. Nuovi Annali delle
VOLUME 99, NUMBER 2
Scienze Naturali e Rendiconto delle Sessioni della
Societa Agraria, e dell’ Accademie delle Scienze
dell’ Istituto di Bologna (2) 2: 256-270, 443-459.
Rondani, C. 1857. Dipterologiae Italicae prodromus.
Vol. 2: Species Italicae ordinis dipterorum in gen-
era characteribus definita, ordinatim collectae,
methodo analitica distinctae, et novis vel minus
cognitis descriptis. Pars prima: Oestridae, Syrpfhi-
dae, Conopidae. 264 pp., 1 fig. Parmae (=Parma).
Ruiz P., FE and Stuardo Oritz, C. 1935. Insectos co-
lectados en las termas de Chillan. Revista Chilena
de Historia Natural 39: 313-322.
Schiner, J. R. 1868. Diptera. In Reise der osterrei-
chischen Fregatte Novara. Zoologia. Vol. 2, Abt.
1, B. 388 pp. Wien.
Shannon, R. C. and Aubertin, D. 1933. Syrphidae.
Diptera of Patagonia and South Chile. Brit. Mus.
(Nat. Hist.) 6: 120-170.
Stuardo Ortiz, C. 1946. Catalogo de los Dipteros de
Chile. 251 pp. Min. Agric. Chile, Santiago.
Tangley, L. 1990. Cataloging Costa Rica’s diversity.
BioScience 40: 633-636.
Thompson, F C. 1972. A contribution to a generic
revision of the Neotropical Milesinae (Diptera:
Syrphidae). Arquivos de Zoologia 23: 73-215.
Thompson, F C. 1981. The flower flies of the West
Indies (Diptera: Syrphidae). Memoirs of the En-
tomological Society of Washington 9, 200 pp. (2
Sep.)
Thompson, E C. 1996. Names: The keys to biodiver-
sity, pp. 199-211. Jn Readka-Kudla, M. L., D. E.
Wilson and E. O. Wilson, eds., Biodiversity II. J.
Henry Press, Washington
Thompson, E C., J. R. Vockeroth and Y. S. Sedman.
1976. Family Syrphidae. A catalogue of the Dip-
tera of the Americas south of the United States
46, 195 pp.
Thompson, F C., J. R. Vockeroth and M. C. D.
Speight. 1982. The Linnaean species of flower
flies (Diptera: Syrphidae). Memoirs of the Ento-
mological Society of Washington 10: 150—165 (30
Sep.)
Townsend, C. H. T. 1895. Contributions to the Dip-
terology of North America.—I. Syrphidae. Trans-
action of the American Entomological Society 22:
33-55.
Townsend, C. H. T. 1897. Diptera from the headwa-
ters of the Gila River. I. Psyche 8: 38-41, 92-94.
Verrall, G. H. 1901. Platypezidae, Pipunculidae, and
Syrphidae of Great Britain. Vol. 8, 691 pp. In his
British Flies. Cambridge.
Walker, F 1852. [Diptera. Continued], pp. 157-252,
pls. v-vi. /n his Diptera. Jn [Saunders, W. W.], ed.,
Insecta Saundersiana. London **1856.”’
Williston, S. W. 1886. Catalogue of the described spe-
cies of South American Syrphidae. Transaction of
the American Entomological Society 14: 308—
324.
237
Williston, S. W. 1891. Fam. Syrphidae, pp. 57-79. In
Godman, FE D. and O. Salvin, eds., Biclogia Cen-
trali-Americana. Zoologia-Insecta-Diptera, vol. 3,
127 pp. (Feb.)
Wilson, E. O. 1985a. The biological diversity crisis:
a challenge to Science. Issues in Science and
Technology 2: 20-29.
Wilson, E. O. 1985b. Time to revive systematics. Sci-
ence 230: 1227.
Wilson, E. O. 1987. The little things that run the
World (The importance and conservation of in-
vertebrates). Conservation Biology 1: 344-346.
Wilson, E. O. 1988. The current status of biological
diversity, pp. 3-18. /n Wilson, E. O. and Peters,
F M., eds., Biodiversity. xiii + 521 pp., National
Academy Press, Washington.
Wilson, E. O. 1992. The diversity of life. 424 pp.
Harvard University Press, Cambridge.
Wulp, E M. van der 1882a. Remarks on certain Amer-
ican Diptera in the Leydon Museum and descrip-
tion of nine new species. Notes from the Leyden
Museum 4: 73-92.
Wulp, E M. van der 1882b. Amerikannsche Diptera.
Tijdschrift voor Entomologie 25: 77—136, pls. 9—
10.
Wulp, E M. van der 1896. Catalogue of the described
Diptera from South Asia. 220 pp. “Dutch Ento-
mological Society,” The Hague.
Wulp, E M. van der 1899. Verbeteringen en Aanvul-
lingen in den Catalogue of the described Diptera
from South Asia. Tijdscrift voor Entomologie 42:
41-57.
Yoon, C. K. 1993. Counting creatures great and small.
Science 260: 620-622.
Zetterstedt, J. W. 1837. Conspectus familiarum, ge-
nerum et specierum dipterorum, in Fauna Insec-
torum Lapponica descriptorum. Isis, Jena 21: 28—
67. (Jan.)
Zetterstedt, J. W. 1838. Sectio Tertia: Diptera, pp.
477-868. In his Insecta Lapponica. vi + 1,140
(pp.). L. Voss, Lipsiae [=Leipzig]. (before Sep.)
Zetterstedt, J. W. 1842—60. Diptera Scandinaviae. Dis-
posita et descripta. Vol. 1, pp. ui—xvi + 1—440
(1842); vol. 2, pp. 441-894 (1843); vol. 3, pp.
895-1280 (1844); vol. 4, pp. 1281-1738 (1845);
vol. 5, pp. 1739-2162 (1846); vol. 6, pp. 2163—
2580 (1847); vol. 7, pp. 2581-2934 (1848); vol. 8,
pp. 2935-3366 (1849); vol. 9, pp. 3367-3710
(1850); vol. 10, pp. 3711-4090 (1851); vol. 11,
pp. v-xii + 4091-4546 (1852); vol. 12: Sup. 3,
pp. v-xx + 4547-4942 (1855); vol. 13: Sup. 4,
pp. v-xvi + 4943-6190 (1859); vol 14, pp. 6191—
6609 (1860). Lundbergiana, Lundae [=Lund].
Zuska, J. and Berg, C. O. 1974. A revision of the
South American genus Tetanoceroides (Diptera:
Sciomyzidae), with notes on colour variations cor-
related with mean temperature. Transaction of the
Royal Entomological Society of London 125:
329-362. (28 Feb.)
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 238-247
A NEW CRYPTIC SPECIES OF TRICHOGRAMMA
(HYMENOPTERA: TRICHOGRAMMATIDAE) FROM THE
MOJAVE DESERT OF CALIFORNIA
AS DETERMINED BY MORPHOLOGICAL,
REPRODUCTIVE AND MOLECULAR DATA
JOHN D. PINTO, RICHARD STOUTHAMER, AND GARY R. PLATNER
(JDP, GRP) Department of Entomology, University of California, Riverside, CA 92521,
U.S.A.; (RS) Department of Entomology, Wageningen Agricultural University, 6700 EH
Wageningen, The Netherlands.
Abstract.—A new species of Trichogramma, T. kaykai, is described from the deserts
of southern California where it is a common egg parasitoid of the lycaenid butterfly
Apodemia mormo. The new species is closely related to T. deion, the most common
Trichogramma in western North America. It is distinguished from 7. deion by morpho-
logical, allozymic and ITS2 sequence differences; the two also appear to be reproductively
incompatible.
Key Words: Trichogramma, new species, allozymes, ITS2 DNA sequences
Collections of parasitized eggs of the ly-
caenid butterfly Apodemia mormo (C. and
R. Felder) on Eriogonum inflatum in the
Mojave Desert in the spring of 1988 were
commonly parasitized by a species of
Trichogramma. This species was originally
considered as a light color form of T. deion
Pinto and Oatman, the most common Trich-
ogramma in western North America (Pinto
et al. 1986). This conclusion was ques-
tioned when typical 7. deion was found at
the same localities also parasitizing A. mor-
mo eggs. Subsequent study showed that this
form differed from T. deion by minor but
consistent morphological traits, and by mo-
lecular differences (allozymes and ITS2 se-
quences). In addition, strains of the two
proved to be reproductively incompatible in
the laboratory. Because this new species
continues to be the subject of various eco-
logical and cytological studies (e.g. Stou-
thamer and Kazmer 1994) it is important
that it receive a formal name. The species
description below is followed by a brief
summary of crossing results, and allozymic
and DNA sequence comparisons. The mo-
lecular data are compared among the new
species, T. deion, T. pretiosum Riley and
the Interior form of T. platneri Nagarkatti.
All taxa are similar morphologically and
occur in sympatry in the Mojave Desert.
SPECIES DESCRIPTION
The description is based on an exami-
nation of material from all localities com-
prising the range of the new species. The
majority of this material consists of wasps
individually mounted on glass slides in
Canada balsam. For body length and color
several specimens of both sexes [reared at
ca. 24° C. and on eggs of Trichoplusia ni
(Hiibner)], and originating from two local-
ities, were critically point dried after being
killed in ethanol and mounted on cards. The
material of 7. deion compared in the diag-
nosis was reared under similar conditions.
VOLUME 99, NUMBER 2
Fig. 1.
Antenna of Trichogramma kaykai male (ventral of left antenna) showing base of flagellum (1500).
Arrows point to positions 1-3 (from right to left) showing presence of a single basiconic peg sensilla at positions
1 and 3 only. In 7. deion, all three positions have a single sensilla.
Unless indicated, quantitative data (means
*+ SD and ranges) are taken from five ran-
domly selected males, all from different lo-
calities. The hind tibial length of these
specimens ranged from 0.16—0.19 mm. The
terminology used in the description follows
Pinto (1992).
Trichogramma kaykai Pinto and
Stouthamer, new species
Trichogramma sp. near deion: Stouthamer
and Kazmer, 1994:317.
Trichogramma sp.: Pinto and Stouthamer,
1994:23 (Table 1.1).
Color sexually dimorphic. Male darker
with mesosoma including coxae orange
brown to yellow brown; pronotum and me-
soscutum typically darkest; metasoma pri-
marily light brown; propodeum and poste-
rior part of scutellum typically lightest, yel-
low orange in color. Female considerably
lighter than male. Most of meso- and me-
tasoma uniformly yellow orange; only pro-
notum, pro- and mesocoxa and ovipositor
suffused with brown; anterior 2—3 metaso-
mal terga lightly suffused with brown in
some specimens.
Body length 0.4—0.5 mm in males, 0.5—
0.6 mm in females.
Forewing 0.28 + 0.02 mm wide, 0.54 +
0.01 as wide as long, setation moderately
dense with 20—29 setae between 4th and
Sth setal tracks, longest fringe setae 0.14-
0.20 maximum forewing width. Hind wing
with 3—4 and 6-8 setae in anterior and pos-
terior tracks, respectively, the latter attain-
ing 0.5—0.6 the distance from hamuli to
apex of wing. Scutellum with anterior pair
of setae short, fine, ca. 0.2 the length of
posterior pair.
Male.—Antenna with flagellum 0.17 +
0.01 mm in length, slightly curved, 6.03 +
0.24 as long as wide, 1.00 + 0.04 as long
as hind tibia, 2.1 + 0.05 as long as scape;
flagellar setae elongate, gradually tapering
to apex, the longest of these setae 3.18 +
0.21 (3.0—3.5) the basal width of flagellum;
basiconic peg sensilla (BPS) relatively
small, only slightly expanded apically, for-
mula 1-0(1)-1-0-1-1 (i.e. position 2 usually
lacking a sensilla as in Fig. 1, also see be-
low for explanation); flagellum lacking un-
socketed setae.
Genital capsule (GC) (Figs. 2, 3) mod-
240
erately broad, 0.34 + 0.02 as wide as long;
sides broadly constricted at level of inter-
volsellar process (IVP); parameres relative-
ly straight, moderately and evenly conver-
gent to apex; apical distance (between apex
of parameres and base of IVP) 0.29 + 0.01
total GC length; apical width (at base of
IVP) 0.63 + 0.2 greatest width of GC; dor-
sal aperature (DA) relatively elongate, nar-
rowing considerably posteriorly, its length
0.63 + 0.03 that of GC; dorsal lamina
(DLA) arising slightly anterior to middle of
GC, moderately notched at base and nar-
rowing directly posterior to notch forming
shoulders which usually do not approach
sides of GC; posterior extension of DLA
relatively elongate, linguiform, its width at
level of IVP subequal to aedeagus width;
DLA length from apex of DA = 0.79 +
0.10 the apical distance and occupying 0.5—
0.6 of this distance, usually extending
slightly beyond apex of volsellae (VS); VS
slightly bowed, occupying 0.4-0.5 the api-
cal distance; IVP narrowly triangular, mod-
erately elongate, occupying 0.3-0.4 the api-
cal distance, its apex usually slightly ante-
rior to that of VS; ventral processes posi-
tioned at base of IVP, slightly protuberant;
ventral ridge relatively short and indistinct,
occupying ca. 0.3 the distance from the
base of the IVP to the base of GC. Aede-
agus length subequal to that of GC and 0.70
+ 0.03 that of the hind tibia; apodemes con-
sisting of ca. 0.5 entire aedeagus length.
Female.—Antenna with funicle segments
subquadrate; usually with 1 BPS on the first
funicular segment; second funicular seg-
ment usually lacking BPS. Ovipositor sub-
equal in length to hind tibia (see below).
Types.—Holotype ¢ and allotype 2
from CALIFORNIA, San Bernardino Co.,
Sheephole Pass, 3 mi. N. on Amboy Rd.;
ex. Apodemia mormo on Eriogonum infla-
tum Torr. & Frém.; iv-30-93; K. Cooper, J.
Pinto & G. Platner, collrs.; in the National
Museum of Natural History, Smithsonian
Institution, Washington, D. C. Eight ¢ and
3 2 paratypes with same data as primary
type deposited in collections of the Cana-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
dian National Collection, Ottawa; Univer-
sity of California, Riverside; University of
California, Berkeley; and The Natural His-
tory Museum, London. All type specimens
are F1-F7 generation individuals from a
culture started with a single female.
Etymology.—The specific name is an ar-
bitrary combination of letters, treated as a
noun in the nominative singular and faithful
to ““KK,”’ the informal epithet applied to
this species in our laboratory for several
years.
Diagnosis.—Trichogramma kaykai is
morphologically most similar to 7. deion
and TJ. pretiosum. All three have similar
genitalic structure and relatively elongate
setae on the male antenna. Males of T. kay-
kai and T. deion are separated from T. pre-
tiosum by the more distinctly sclerotized
posterior extension of the dorsal lamina and
the BPS formula on the antenna. Unlike T.
kaykai and T. deion, T. pretiosum has a pair
of BPS at the second and third positions.
Differences in this trait and male genitalia
in T. deion and T. pretiosum are summa-
rized in Pinto et al. (1986).
Trichogramma kaykai is separated from
T. deion by color, minor but consistent dif-
ferences in the male genitalia, BPS formula,
and ovipositor length. Color in T. deion is
not obviously sexually dimorphic as it is in
T. kaykai. In T. deion the mesoscutum and
metasoma of both sexes are typically
brownish. Males of T. kaykai are similar but
usually a lighter brown; females, however,
are almost uniformly yellow or yellow or-
ange and are easily separated from T. deion
on this basis.
In most Trichogramma there are one or
two basiconic peg sensilla (BPS) at each of
the first three positions of the male flagel-
lum. The presence of these sensilla are rel-
atively stable within species, although they
may be reduced or completely absent at
these positions in abnormally small individ-
uals. T. deion almost always has a single
BPS at the first three positions. In T. kaykai
a single sensilla is present at positions 1 and
3 but usually not at position 2 (Fig. 1). This
VOLUME 99, NUMBER 2
DLA
DA
2 3
Figs. 2—4.
241
Genital capsule of male (posterior end above). 2, Trichogramma kaykai (dorsal view), DLA =
dorsal lamina, DA = dorsal aperature, VS = volsella. 3, T. kaykai (ventral view). 4, T. deion (dorsal view).
Note in 7. kaykai the DA is narrower apically, and the DLA is narrower at its base (immediately posterior to
notch) and extends beyond the apex of the VS.
feature is variable, however. Whereas the
formula in JT. deion is relatively stable,
about 35% of 135 non-sib males of T. kay-
kai examined had a sensilla at this position.
Females of T. kaykai can also usually be
separated from 7. deion by the absence of
a BPS at the equivalent antennal position,
i.e., the apex of the second funicular seg-
ment.
Most males of T. kaykai can be separated
from T. deion by genitalia structure (cf
Figs. 2, 4). In T. kaykai the genital capsule
is narrower apically, the dorsal lamina is
narrower immediately posterior to its basal
notch, and, most importantly, the dorsal
aperature is somewhat longer and narrows
considerably at its posterior end. Correlated
with this difference, the length of the dorsal
lamina, measured from the posterior border
of the dorsal aperature to its apex, is less in
T. kaykai and averages 0.79 the apical dis-
tance; in T. deion this ratio averages 0.93
the apical distance. Also, in T. kaykai the
lamina typically extends to a level slightly
posterior to the apex of the volsellae; it usu-
ally does not extend that far in T. deion.
Females of T. kaykai are most easily dis-
tinguished by color. However, the longer
242
ovipositor also helps distinguish it from T.
deion. Samples of 10 females of each spe-
cies from localities where the two were
sympatric were compared. All originated
from different host eggs and had a hind tib-
ial length ranging from 0.19—0.21 mm. In
T. kaykai, the ratio of ovipositor to hind tib-
ial length was 1.00 + 0.03; in T. deion this
ratio was 0.93 + 0.04. Although the range
of variation does overlap, this feature used
with the significantly lighter coloration and
absence of a BCP on the second funicular
segment should provide straightforward
separation of females.
Geographic distribution.—Trichogram-
ma kaykai has only been collected in south-
ern California -in the Mojave Desert and at
the northern limits of the Sonoran Desert.
Hosts.—The primary host of this species
is the egg of Apodemia mormo (Lycaeni-
dae) laid on Eriogonum inflatum. This ly-
caenid has been divided into several sub-
species (Miller and Brown 1981). All re-
cords of T. kaykai are from A. m. deserti
Barnes and McDunnough. Trichogramma
kaykai also has been taken a single time
(Pinyon Mt., Kern Co.) on the egg of an-
other lycaenid, Jcaricia lupini (Boisduval),
on Eriogonum fasciculatum Bentham.
Material examined.—401 specimens of
both sexes. The material examined includes
specimens emerging from field collected
host eggs and from cultures initiated with
these parentals.
Records.—Except for the single record
from Pinyon Mt. (see Hosts), all following
collections originated from the eggs of Apo-
demia mormo on Eriogonum inflatum.
UNITED STATES. California. Kern Co.
El Paso Mts. (Bickel Camp); vii-29-96; R.
Stouthamer. Pinyon Mt.; vi-4-87; G. Pratt.
Walker Pass, 2 mi. E.; iv-19-89; G. Pratt.
Last Chance Cyn.; v-14-88; D. Kazmer, R.
Stouthamer; also numerous collections by
several collectors from iv-vi-95/96. Rands-
burg; v-14-88; D. Kazmer, R. Stouthamer.
Riverside Co. Dillon Rd., several collec-
tions between immediately east of Indio
and Fun Valley (all thelytokous); iv-26-95;
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
J. Pinto, R. Stouthamer. San Bernardino
Co. Barstow, ca. 10 mi. NE., and ca. 8 mi.
E.; v-29-96, v-15-96; L. Bolijn, B. Deijkers.
Beacon Station; vi-6-96; L. Bolijn, B.
Deijkers. Danby; iv-11-88, iv-30-88, iv-27-
95; G. Pratt/D. Kazmer/R. Stouthamer.
Joshua Tree; vi-15-96; L. Bolijn, B. Deijk-
ers. Hwy. 66 between Bagdad and Siberia;
iv-27-95; R. Stouthamer. Interstate 15, be-
tween Victorville and Barstow; v-29-96; L.
Bolijn, B. Deijkers. Kramer Hills; v-14-88,
iv-24-95; D. Kazmer, R. Stouthamer/J. Ben-
nett, R. Stouthamer. Lucerne Valley, ca. 12
mi. N.; v-29-96; L. Bolijn, B. Deijkers.
Sheephole Pass area; v-14-88, iv-27-95; D.
Kazmer, R. Stouthamer. Sheephole Pass, 3
mi. N. (on Amboy Rd.) (type locality);
iv-30-93; K. Cooper, J. Pinto, G. Platner.
Yucca Valley; vi-1-96; L. Bolijn, B. Deijk-
ers.
Notes.—Both thelytokous and arrheno-
tokous populations of 7. kaykai have been
collected. As in many species of Tricho-
gramma, thelytoky in this species is caused
by Wolbachia infection and can be cured
with antibiotic treatment (Stouthamer et al.
1990). Populations with both modes of re-
production occur together in the Mojave
Desert. The few collections from the So-
noran Desert are all thelytokous.
Unlike 7. deion which has very broad
host and geographic ranges in western
North America (Pinto et al. 1986), T. kaykai
has been retrieved almost totally from eggs
of Apodemia mormo in the deserts of south-
ern California. Both species are known to
occur together on Apodemia at several lo-
calities, although 7. kaykai is more com-
mon and appears to be the dominant egg
parasite of this butterfly in the Mojave Des-
ert. Of a sample of 256 parasitized A. mor-
mo eggs collected at several sites of sym-
patry during the spring of 1995, 212 or
82.9% were attacked by T. kaykai; only 38
or 14.8% were parasitized by T. deion. The
remainder (2.3%) were attacked by a third
and undescribed species. In a few cases, T.
deion and T. kaykai emerged from the same
egg.
VOLUME 99, NUMBER 2
The eggs of A. mormo are relatively large
and several Trichogramma typically
emerge from a single egg. In a sample of
33 eggs collected in 1988 from several lo-
calities the average number of T. kaykai
emerging from each was 4.6 + 1.2 (range
= 1-7). Most of the progeny were female;
most parasitized eggs result in a single male
and 3—4 females. The average number of
males emerging from this sample (exclud-
ing eggs resulting in thelytokous wasps)
was 1.19 + 0.75 (range = O-3, n = 31).
Collections of other species of host at
sites where T. kaykai occurs are minimal.
The only such collection was made on 14
May 1988 at Last Chance Canyon in Kern
County. The eight parasitized eggs of A.
mormo collected on Eriogonum inflatum
were attacked by both 7. deion (5) and T.
kaykai (3). However, only T. deion emerged
from 23 parasitized eggs of an undeter-
mined Pieridae collected from a species of
Stanleya (Brassicaceae).
MOLECULAR DATA
Allozymes
An earlier paper compared allozymes in
T. pretiosum and T. deion (Pinto et al.
1993) and reported consistent allelic differ-
ences between these close relatives. For this
study we examined ten loci in four cultures
of T. kaykai, and compared them with one
culture of the interior form of T. platneri
Nagarkatti, two cultures of 7. pretiosum
and seven of 7. deion. All exemplars were
run concurrently. The cultures chosen of the
latter three species represented most of the
known allelic diversity in Trichogramma at
the loci examined. The enzyme systems
used are as follows: Aconitase (4.2.1.3),
Acon; acid phosphatase (3.1.3.2), Acp-II;
fumarase (4.2.1.2), Fum; a-glycerolphos-
phate dehydrogenase (1.1.1.8), aGpd-IT;
glucose-phosphate isomerase (5.3.1.9), Gpi;
glucose-6-phosphate dehydrogenase (1.1.1.49),
G6pd; isocitrate dehydrogenase (1.1.1.42),
Idh; malate dehydrogenase (1.1.1.37), Mdh-
IT; malic enzyme (1.1.1.40), Me; and phos-
243
phoglucomutase (2.7.5.1), Pgm. These loci,
among others, were also compared between
T. pretiosum and T. deion in our earlier
study.
The four T. kaykai cultures examined
were from Walker Pass (KWPA), Last
Chance Cyn. (KLC187), and between Bag-
dad and Siberia (KRB85), three Mojave
Desert localities, and Dillon Rd. (N. of In-
dio) (KAW73), a Sonoran Desert site. The
cultures of 7. deion examined represent
much of the range of the species in western
North America. The origin of the compar-
ison cultures were as follows: T. pretiosum
— Riverside, CA (PRV4) and Wyndham,
Australia (PAWD). Interior form of T. plat-
neri—Mesquite, NV (IMSQ). 7. deion —
Riverside, CA (DRV4); Seven Pines, CA
(DSVP); Covelo, CA (DCLO); Portal, AZ
(DPTL); Granite Gap (Hidalgo Co.), NM
(DGGP); Miles City, MT (DMCT); Paul’s
Place (Kern Co.), CA (DPPL); and Last
Chance Cyn (DLC1). All but two of these
comparison cultures (DPPL and DLC1)
also were examined in our previous paper
(Pinto et al. 1993). The culture IMSQ was
originally assigned to the interior form of
T. platneri (Pinto et al. 1992). This form
represents a new species and will be de-
scribed in the near future. Until then we
continue to refer to it as before.
All cultures compared represented iso-
female lines. One culture of 7. kaykai
(KAW73) was originally thelytokous; all
others were arrhenotokous. In the latter
case, the female used to initiate a culture
had mated with a brother.
Electrophoretic analysis followed meth-
ods in our earlier studies (Pinto et al. 1992,
1993) and were originally detailed by Kaz-
mer (1991). Briefly, two females per culture
were individually analyzed at each locus by
isoelectric focusing in one to two layers of
cellulose acetate membranes using a single
blend of carrier ampholytes (8% pH 4-—6.5
and 2% pH 3-10 pharmalytes) (Sigma
Chemical, St. Louis, MO) and an effective
gel length of 4.5 cm. Each culture repre-
sented an isofemale line initiated from a
244
ables:
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Allelic comparison of Trichogramma kaykai with closely related species at eight loci.*”
Loci and Alleles
Taxon Culture* Acpll Fum Gpi aGpdll G6pd Idh Mdhil Pgm
Interior form IMSQ D G B A A A G E
pretiosum PRV4 D B B A B B A D
PAWD D B B A B B E E
deion DRV4 € B B B A A ‘S B
DSVP D B A A A B S €
DCLO (C B B A A B Ec B
DPTL D B B A A A B g
DGGP D B B A A B ‘S B/C
DMCT D B B B A A C A
DPPL Cc B B A A B Cc B
DECIC — — B — A — Cc —
kaykai KWPA B B (@ A C/F B A B
KRB85 D B C A D/E B A B
KLC187 D € D A D/E B A B
KAW73 D B C A CE B A B
4 Relative distances among electromorphs for the three loci distinguishing 7. kaykai [distances for others given
in Pinto et al. 1993)], based on ratio of distance between cathode and homomeric band to entire gel length, as
follows (alleles in alphabetical order): Gpi (0.18, 0.46, 0.62, 0.67), G6pd (0.03, 0.11, 0.19, 0.23, 0.27, 0.32),
MadhlII (0.31, 0.38, 0.47, 0.56).
> All females examined were homozygous at all loci. Those entries showing two alleles indicate the two females
examined were homozygous for different alleles.
© See text for geographic origin of cultures.
4DLCl1, a T. deion collection sympatric with the 7. kaykai KLC187, was only examined at the three loci
distinguishing the two species.
single parasitized host egg collected in the
field and maintained in the laboratory for
several generations on eggs of Trichoplusia
ni. BIOSYS-1 (Swofford and Selander
1989, release 1.7) was used to analyze data
using individual genotypes as input.
Results.—Two loci (Acon and Me) were
fixed in all samples examined. The remain-
ing eight were polymorphic and are com-
pared for all cultures in Table 1. The four
T. kaykai cultures differed from all heter-
ospecifics at two loci, Gpi and G6pd. One
of these loci, G6pd, is the only one provid-
ing complete separation of 7. deion and T.
pretiosum (Pinto et al. 1993). The three al-
lozymic differences between T. kaykai and
T. deion also were found in one pair of
sympatric collections (from Last Chance
Cyn., Kern Co., CA). Cluster analysis based
on the data in Table 1, using UPGMA clus-
tering of Nei genetic distances, recognized
three groups, all consistent with species
identity. IMSQ joined closest to the pre-
tiosum cultures. Mean Nei genetic distances
between 7. kaykai and T. deion, and be-
tween 7. kaykai and T. pretiosum were
0.697 + 0.24 (0.379-1.178, n = 27) and
0.490 + 0.13 (0.311—0.668, n = 8), respec-
tively. The mean intraspecific distance for
T. kaykai was 0.218 + 0.14 (0.027—0.460,
n = 6). The interspecific distances are con-
siderably greater than those reported in our
earlier studies of closely related Tricho-
gramma and are certainly exaggerated. This
is because cultures of both T. deion and T.
pretiosum were chosen for allelic diversity
to insure detection of any differences that
occurred in the new species.
The presence of at least two alleles of
G6pd in the cultures of T. kaykai (Table 1)
requires further investigation. Although this
could be explained if the female initiating
each culture was heterozygous, it is also
possible, and perhaps more likely, that two
loci for this enzyme are involved.
VOLUME 99, NUMBER 2
245
Table 2. The size of the ITS2 gene and size of the restriction fragments generated by restriction enzymes
MSE1 and ECORI.
MSE} restr. ECOR| restr
Size ITS2
Taxon Culture* (bp) fragments (bp) fragment (bp)
T. deion DSVP 398 294, 61, 43 398
DCLO 402 296, 63, 43 402
DLC1 406 300, 65, 41 406
T. kaykai KLC187 470 271, 199 470
KSH1 463 263, 200 463
T. pretiosum PRV4 409 409 409
PIRV 413 413 413
Interior form IMSQ SIS 409, 81, 25 325, 190
4 See text for geographic origin of cultures.
ITS2 DNA Sequences
The ribosomal nuclear genes (rRNA) are
among the several genetic sequences pro-
posed for distinguishing closely related spe-
cies in insects (Hoy 1994). The ITS2 se-
quence, or Internal Transcribed Spacer re-
gion, is positioned between the 5.8S and
28S coding region of the rRNA gene. This
sequence shows considerable promise in
separating closely related species of Trich-
ogramma (van Kan, et al. 1996). In this
study we compared the complete ITS2 se-
quences of two T. kaykai cultures, three T.
deion and two T. pretiosum cultures, and
one culture of the interior race of T. platneri
(see Table 2). Each culture represented an
isofemale line initiated with a single para-
sitized host egg collected in the field and
maintained for a variable number of gen-
erations on eggs of either Trichoplusia ni,
Mamestra brassicae L. or Ephestia kuehn-
iella (Zeller). Differences in sequence and
size of the ITS2 gene were determined. Re-
striction enzymes were used to find char-
acteristic differences among species. Cul-
tures studied are indicated in Table 2. Sev-
eral of these were also used for allozymes
(Table 1). Those examined for ITS2 only
include the following: T. kaykai—Last
Chance Cyn., CA (KLC187); Sheephole
Pass area (San Bernardino Co.) (KSH1). T.
pretiosum—Irvine, CA (PIRV). Cultures
included both allopatric and sympatric
(DLC1 & KLC187) representatives of T.
deion and T. kaykai.
The method for determining the ITS2 se-
quence, briefly, is as follows: One to three
wasps, preserved in absolute ethanol, were
ground in 50-150 pl 5% Chelex and 3 ul
proteinase K (20 mg/ml) and incubated for
at least 2 hrs. at 56°C followed by 10 min
at 95°C. The wasps were first shaken in 1
ml TAE for 1 hr prior to grinding in Chelex.
PCR was performed in 50 ul reaction vol-
umes using a Hybaid thermocycler, 5 pl
DNA template, 5 pl PCR-buffer, 1 pl d-
NTP’s (each in a 10mM concentration), 0.6
wl forward and reverse primer (10ng), 0.1
wl SuperTth polymerase enzyme (5 units/
ul) from Spaero-Q; and 38 wl sterile dis-
tilled water. The ITS-2 region was ampli-
fied using the following primers; forward:
5’-TGTGAACTGCAGGACACATG-3’; re-
verse: 5’°-AATGCTTAAATTTAGGGGGTA-
3’. The PCR cycling program was 3 min.
95°C; 45. sec. at, 533 and’ 45sec... ato/2°C
with 3 min. at 72°C after the last cycle. The
machine was set to tube control. PCR prod-
ucts of about 550 bp were electrophoresed
and excised from the agarose gel. They
were then frozen and freeze-squeezed. The
liquid phase was alcohol precipitated,
washed and ligated into a T-tailed vector
(Amersham Life Science) and amplified in
E. coli cells. Escherichia coli colonies con-
taining an insert of the correct size were
checked by PCR using the primers men-
tioned above and were subsequently se-
quenced using an automatic sequencer (373
DNA Sequencer Stretch, Applied Biosys-
246
tems using a Prism Ready Reaction
DyeDeoxy6é Terminator Cycle sequence
kit). The size of digestion products of the
ITS2 gene using different restriction en-
zymes was determined. Characteristic dif-
ferences among the species were found us-
ing ECORI and MSE1.
Results.—The sequences of the ITS2
genes have been deposited in the EMBL,
GenBank and DDBJ Nucleotide Sequence
Databases. Accession numbers for the spe-
cies and cultures indicated in Table 2 are as
follows: T. deion—U76223 (DCLO),
U76224 (DLC1), U76225 (DSVP); T. pre-
tiosum—U76226 (PRV4), U76227 (PIRV);
T. kaykai—U76228 (KSH1), U76229
(KLC1); Interior form of T. platneri—
U76230 (IMSQ).
The size of the 7. kaykai ITS2 gene is
consistently larger than that of either T.
deion and T. pretiosum. The size of the
ITS2 gene in order from large to small is:
Interior form of T. platneri > T. kaykai >
(T. deion, T. pretiosum). The consistent and
characteristic differences found in the ITS2
sequences in these four species are reflected
in differences in restriction length frag-
ments when the ITS2 gene is restricted with
the enzymes MSE! and ECORI as shown
in Table 2. The interior form of T. platneri
differs from the other species in the size of
its ITS2 gene (515 bp) and in the presence
of the restriction site for ECORI. Tricho-
gramma kaykai differs from both T. deion
and T. pretiosum in the size of its ITS2
gene (470 bp vs 400—410 bp). Also, the
ITS2 gene of T. kaykai is cut in two large
fragments by MSE1 (270 and 200 bp)
whereas T. deion is cut into three fragments
with the largest ca. 300 bp; T. pretiosum is
not restricted by this enzyme. As with al-
lozymes, the differences between T. deion
and T. kaykai occur in both allopatric and
sympatric collections.
REPRODUCTIVE DATA
Reproductive compatibility has frequent-
ly been used to support species hypotheses
in Trichogramma (Pinto and Stouthamer
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1994). In this study, four cultures of T. kay-
kai were crossed (at 25°C) with a culture of
T. deion from Dillon Rd., near Indio, CA
(DAW6) and among themselves. The T.
kaykai cultures used included two from
Danby, CA (KDA22, KDA23), and two
from Last Chance Cyn., CA (KLC18,
KLC21). Crossing procedures employed
closely followed those detailed in Pinto et
al. (1991). All crosses are based on individ-
ual pairings. All combinations of hetero-
gamic crosses were performed concurrently
with homogamic controls, resulting in a to-
tal of 20 crosses. All crosses included 20
replicates in each direction. For determin-
ing relative compatibility between cultures
the mean sex ratio (MSR) was calculated as
the percentage of female progeny. The rel-
ative compatibility of an interculture cross
(A X B) is expressed as two percentages:
100% X MSR (A female X B male)/MSR
(A female X A male); and the same based
on the reciprocal.
Results.—The crosses between cultures
of T. kaykai and T. deion were completely
incompatible. Female production is the ev-
idence for reproductive compatibility in ar-
rhenotokous Hymenoptera and not a single
female was produced in any of the four
heterospecific trials. The mean relative
compatibility among the 12 homospecific
crosses of T. kaykai cultures was 78.0 and
ranged from 49.0 to 97.5. The least com-
patible cross (49.0) was between KDA22
females and KLC21 males. The reciprocal
cross was considerably stronger (83.1).
There was no evidence that the sympatric
cultures (mean compatibility = 74.2, n =
4) were more compatible than the allopa-
triates (mean compatibility = 79.8, n = 8).
In fact, the highest levels (92.6, 97.5) were
among allopatric cultures.
CONCLUDING REMARKS
Morphologically similar species of
Trichogramma as exemplified by T. kaykai
and T. deion are apparently quite common
(Pinto and Stouthamer 1994). Yet, we feel
that in this genus such cryptic species
VOLUME 99, NUMBER 2
should be described only after putative
morphological differences are shown to be
geographically stable and, ideally, found to
correlate with other character sources. This
requires extensive collecting of all forms
involved and an attempt to delineate at least
rough geographic distributions. One goal of
such work should be the identification of
areas of sympatry since it is at such local-
ities that the stability of character differ-
ences can be most rigorously tested for. In
the case of T. kaykai, we have shown that
this species has only minor morphological
differences from 7. deion but that these are
consistent at several localities including
those where the two occur together. Differ-
ences in allozymes and the ITS2 sequence,
as well as crossing incompatibility, give
greater confidence that species recognition
is warranted. Thus far we have no evidence
of gene flow between these two species.
However, a focus of future work should in-
clude additional crossing and molecular
studies in zones of sympatry. Our efforts in
both areas are preliminary.
ACKNOWLEDGMENTS
We thank Gordon Pratt, University of
California, Riverside, for his contribution to
this study. He was the first to collect the
new species and also provided valuable in-
formation on its host, A. mormo. We also
thank EF J. PR M. van Kan for determining
the ITS2 sequences reported in this study,
and L. Bolijn and B. Deijkers for several of
the collections made in the Mojave Desert
in the spring of 1996.
LITERATURE CITED
Kazmer, D. J. 1991. Isoelectric focusing procedures
for the analysis of allozymic variation in minute
arthropods. Annals of the Entomological Society
of America 84:332—339.
Hoy, M. 1994. Insect molecular genetics. An intro-
duction to principles and applications. Academic
Press Inc.
Miller, L. D. and EK M. Brown. 1981. A catalogue/
247
checklist of the butterflies of America north of
Mexico. The Lepidopterists’ Society Memoir No.
2, 280 pp.
Pinto, J. D. 1992. Novel taxa of Trichogramma from
the New World tropics and Australia (Hymenop-
tera: Trichogrammatidae). Journal of the New
York Entomological Society 100:621—633
Pinto, J. D. and R. Stouthamer. 1994. Systematics of
the Trichogrammatidae with emphasis on Tricho-
gramma, Chapter 1, pp. 1-36. In Wajnberg, E. and
S. A. Hassan, eds., Biological control with egg
parasitoids. CAB International, Wallingford.
Pinto, J. D., E. R. Oatman and G. R. Platner. 1986.
Trichogramma pretiosum and a new cryptic spe-
cies occurring sympatrically in southwestern
North America (Hymenoptera: Trichogrammati-
dae). Annals of the Entomological Society of
America 79:1019—1028.
Pinto, J. D., R. Stouthamer, G. R. Platner and E. R.
Oatman. 1991. Variation in reproductive compat-
ibility in Trichogramma and its taxonomic signif-
icance (Hymenoptera: Trichogrammatidae). An-
nals of the Entomological Society of America 84:
37-46.
Pinto, J. D., D. J. Kazmer, G. R. Platner and C. A.
Sassaman. 1992. Taxonomy of the Trichogram-
ma minutum complex (Hymenoptera: Trichogram-
matidae): allozymic variation and its relationship
to reproductive and geographic data. Annals of the
Entomological Society of America 85:413—422.
Pinto, J. D., G. R. Platner and C. A. Sassaman. 1993.
Electrophoretic study of two closely related spe-
cies of North American Trichogramma: T. pre-
tiosum and T. deion (Hymenoptera: Trichogram-
matidae). Annals of the Entomological Society of
America 86:702—709.
Stouthamer, R., R. EF Luck and W. D. Hamilton. 1990.
Antibiotics cause parthenogenetic Trichogramma
to revert to sex. Proceedings of the National Acad-
emy of Sciences, USA 87:2424—-2427.
Stouthamer, R., and D. J. Kazmer. 1994. Cytogenetics
of microbe-associated parthenogenesis and its
consequences for gene flow in Trichogramma
wasps. Heredity 73:317—327.
Swofford, D. L. and R. B. Selander. 1989. BIOSYS-1.
A computer program for the analysis of allelic
variation in population genetics and biochemical
systematics. Release 1.7. Urbana, Illinois.
van Kan, FJ.P.M., I. M. M. S. Silva, M. Schilthuizen,
J. D. Pinto and R. Stouthamer. 1996. Use of
DNA-based methods for the identification of min-
ute wasps of the genus Trichogramma. Proceed-
ings of the section Experimental and Applied En-
tomology of the Netherlands Entomological So-
ciety. 7:233—237.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 248-256
A NEW SPECIES OF TRYPANARESTA HERING (DIPTERA: TEPHRITIDAE)
FROM PATAGONIA, A POTENTIAL AGENT FOR BIOLOGICAL CONTROL
OF SNAKEWEEDS (GUTIERREZIA SPP.) IN THE UNITED STATES
DANIEL E. GANDOLFO AND ALLEN L. NORRBOM
(DEG) USDA-ARS, South American Biological Control Laboratory, Bolivar 1559
(1686) Hurlingham, Buenos Aires province, Argentina; (ALN) USDA-ARS-PSI, System-
atic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Ag-
riculture, c/o National Museum of Natural History, MRC 168, Washington, D.C. 20560,
U.S.A.
Abstract.—Trypanaresta valdesiana, n. sp., is described. Illustrations of the egg, third
instar larvae, puparia and adults are provided. Larvae develop inside the buds of Gutier-
rezia solbrigii Cabrera and Gutierrezia mandonii (Schultz Bipontinus) Solbrig in Pata-
gonia, Argentina. No seeds are produced by infested capitula. Larvae overwinter inside
the dry infested buds and adults emerge in spring. Two Hymenoptera, Torymoides sulcius
(Walker) (Torymidae) and Epicatolaccus strobeliae Blanchard (Pteromalidae) are endo-
parasitoids of larvae and pupae. Trypanaresta valdesiana is currently being studied as a
potential agent for biocontrol of snakeweeds, Gutierrezia spp., in the United States.
Key Words:
Trypanaresta, Tephritidae, Gutierrezia, snakeweed, Asteraceae, weed bio-
control, taxonomy, immature stages
The genus Gutierrezia (Asteraceae: As-
tereae) originated in North America and
several species are endemic to the south-
western United States and northern Mexico
(Solbrig 1960, Lane 1985). Two perennial
species, G. sarothrae (Pursh) Britton &
Rusby (broom snakeweed) and G. micro-
cephala (D.C.) Gray (threadleaf snake-
weed), and two annuals, G. texana (D.C.)
Gray and G. spharocephala Gray, are wide-
spread and serious weeds of the semiarid
rangelands of the southwestern United
States. Twelve species of Gutierrezia are
endemic to South America, seven from Ar-
gentina and five from Chile (Solbrig 1966,
Cabrera 1971). All South American species
are perennials and have low or no economic
impact.
In the U.S.A. losses due to snakeweed
have been estimated to be at least $34 mil-
lion per year (McGinty and Welch 1987,
Cordo and DeLoach 1992). Although
chemical control is possible, two major
problems have almost completely precluded
its use: 1) the low economic return of the
infested rangelands, and 2) the unpredicta-
bility of natural fluctuations in snakeweed
populations (DeLoach 1981, Torell et al.
1990). Biological control as an alternative
against native weeds using natural enemies
of the South American species has been
proposed and discussed by DeLoach
(1981). Cordo and DeLoach (1992) listed
the natural enemies of the Argentine Gu-
tierrezia species and discussed climatic and
ecological similarities of the troublesome
Gutierrezia species and the Argentine spe-
Gies:
The purpose of this paper is to describe
a new species of Trypanaresta that is cur-
VOLUME 99, NUMBER 2
rently being studied at the South American
Biological Control Laboratory, ARS-
USDA, Hurlingham, Argentina (SABCL)
as a potential agent for the biological con-
trol of snakeweeds in the southwestern
United States.
MATERIALS AND METHODS
Adults were collected or reared from
samples taken in Patagonia, Argentina,
from 1993-95. Most of them were, reared
from mature larvae or pupae in samples of
flower heads of Gutierrezia solbrigii Ca-
brera collected near Puerto Piramide, Chu-
but province, Argentina. The morphological
terminology used for adults follows Foote
et al. (1993), for female genitalia Norrbom
and Kim (1988), and for larvae Teskey
(1981). Measurements were taken from 10
specimens of each sex, as described by Jen-
kins and Turner (1989). The lengths of syn-
tergosternite 7, eversible membrane, and
aculeus were measured ventrally on dis-
sected specimens mounted on a microscope
slide (n = 5). Female genitalia were pre-
pared using the technique described in
Foote et al. (1993). Larval spiracles and
mouthparts were prepared as described by
Steck and Wharton (1986).
The following acronyms are used for
specimen depositories: MACN, Museo Ar-
gentino de Ciencias Naturales, Buenos Ai-
res, Argentina; SABCL, South American
Biological Control Laboratory, Hurlingh-
am, Argentina; USNM, National Museum
of Natural History, Smithsonian Institution,
Washington D.C., U.S.A. Gutierrezia spp.
were identified by A. Cabrera of the Insti-
tuto Darwinion, San Isidro, Argentina. Par-
asitoids were identified by L. De Santis
(Museo de Ciencias Naturales, La Plata, Ar-
gentina). Voucher specimens for plants and
parasitoids are deposited at the SABCL.
TAXONOMY
The genus Trypanaresta was proposed
by Hering (1940) for a group of species
similar to Trupanea in having a subapical
stellate mark in the wing pattern, but with
249
2 pairs of scutellar setae (the apical at most
half as long as the basal), frons setulose, 2—
3 frontal and 2 orbital setae. Foote (1967)
listed nine species in Trypanaresta, but, in
addition, all the Neotropical species previ-
ously placed in Goniurella (Foote 1980)
and most South American species previ-
ously classified as Tephritis (Foote 1967,
1980; Frias 1988) belong in this genus
(Norrbom, in prep.). The wing pattern is
more extensive in some species than in
those originally included by Hering, and the
apical scutellar seta is actually minute or
absent in a few species, but Norrbom
(1993: 205) noted that all species of Try-
panaresta lack a pair of small but outstand-
ing dorsal preapical setulae on the hind fe-
mur. This character diagnoses Trypanaresta
and the closely related genus Plaumanni-
myia from Trupanea, Euaresta, and other
similar Neotropical genera of Tephritini.
Trypanaresta valdesiana Gandolfo and
Norrbom, new species
(Figs. 1-3)
Type material.—Holotype: 6 (MACN)
ARGENTINA: Chubut: near Puerto Pir-
amide, 25-III-1995, Gandolfo & Calcaterra,
reared from larva in flower head of Gutier-
rezia solbrigii (197097). Paratypes: same
data as holotype, 6 6, 5 2 (USNM,
MACN, SABCL); same except 10-I-1994,
D. Gandolfo (153800), 12 (USNM); same
except 24-I-1994, D. Gandolfo, 1
(SABCL); same locality, 9-IX-1994, as
overwintering larva in dry capitulum of
Gutierrezia solbrigii, Gandolfo (174431), 3
3,2 2 (SABCL); same locality, 25-I-1994,
as larvae in capitula of Gutierrezia solbri-
gii, D. Gandolfo, 12 (SABCL); ARGEN-
TINA: Chubut: 11 km. N. Puerto Madryn,
27-HI-95, Gandolfo & Calcaterra (197429),
12 (SABCL); Puerto Madryn, 27-III-95,
Gandolfo & Calcaterra, reared from larva
in flower head of Gutierrezia_ solbrigii
(197468), 12 (USNM); 44 km. SW of Pun-
ta Norte, 20-XII-93, D. Gandolfo, reared
from capitulum of Gutierrezia_ solbrigii
(149792), 1 6, 1 2 (USNM); Rio Negro:
Fig. 1. Trypanaresta valdesiana (A) head, lateral
view; (B) wing; (C) wing apex with medial ray in cell
T>,3 Incomplete; (D) same, with medial ray absent.
San Antonio Oeste, 8-XII-93, D. Gandolfo,
reared from capitula of Gutierrezia solbrigii
(149010), 1¢ (USNM). Additional speci-
mens examined: 12 (SABCL) ARGENTI-
NA, La Pampa, Lihue Calel, 13-I-1995,
Gandolfo & Velazquez, reared from capit-
ulum of G. mandonii (1785); 22 164
(SABCL) ARGENTINA, Rio Negro, 17-I-
1995, Gandolfo & Velazquez, 4km. W Ra-
mos Mexia, reared from capitula of G. sol-
brigii; 12 (SABCL) ARGENTINA, Neu-
quén, Arroyito 16-I-1996, Gandolfo & Ve-
lazquez, reared from larva in flower head
of Gutierrezia solbrigii.
Diagnosis.—T. valdesiana can be distin-
guished from all other species of Trypana-
resta by the following combination of char-
acters: apical scutellar seta present; male
foreleg unmodified; wing pattern (Fig. 1b)
uniformly dark brown; pterostigma con-
nected with r-m by solid dark brown band;
discal cell without subapical rays, hyaline
except for margin of the broad diagonal
band, which extends slightly posterior to
vein M between crossveins r-m and dm-cu;
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and wing basad of pterostigma and at least
basal % of cell cu, hyaline.
Description.—Adult: Body length 3.31-
4.13 mm, female barely larger than male.
Most setae yellow brown to brown. Head
(Fig. la): barely higher (0.65—0.91 mm)
than long (0.48—0.70 mm). Generally yel-
low, but frons yellow to brown. Frons at
vertex wider (0.49—0.72 mm) than long
(0.36—0.50 mm), slightly narrowed to an-
terior margin (0.45—0.65 mm). Anterior half
of frons with a few whitish setulae smaller
than numerous fronto-orbital setulae (these
frontal setulae rub off easily and may not
be present in poorly preserved specimens).
2 frontal and 2 orbital setae, posterior or-
bital smaller, whitish and inflated. Ocellar
tubercle blackish, with white setulae. An-
tenna testaceous-yellow, first flagellomere
about as long as wide. Head setae equally
long in both sexes. Outer vertical, post-
ocellar and postocular setae whitish. Tho-
rax: Ground color of scutum black, of scu-
tellum yellow to pale brown. Mesonotum
covered by bright yellow microtrichia ex-
cept anterior central area of scutum with
pale gray microtrichia. Setulae white to yel-
low, inflated, relatively long (twice as long
as in T. thomsoni). Scutellum with 2 pairs
of setae, apical pair about % as long as ba-
sal. Anepisternal, katepisternal, anepimeral,
and posterior notopleural setae white to yel-
low (concolorous with setulae), other tho-
racic setae yellow brown to brown. Wing:
3.0-3.5 mm long and 1.1—1.6 mm wide.
Wing pattern (Fig. 1b) uniformly dark
brown. Pterostigma connected to stellate
mark by broad, solid band, without hyaline
spots; base of band somewhat truncate, not
extended to base of R,,; nor with hyaline
indentation in cell r,,;. Discal cell totally
hyaline except for margin of broad diagonal
band, which reaches slightly beyond vein M
between crossveins r-m and dm-cu. Dark
ray over dm-cu and 2 dark rays crossing
cell m all reaching posterior wing margin.
Center of stellate mark without hyaline
spots; basal % of cell r,,; dark except for
hyaline spot anterior to dm-cu, touching
VOLUME 99, NUMBER 2
vein M, but never extended to R,,,;. Cell r,
with 3 hyaline marginal marks: the largest,
immediately distad of pterostigma, oblique
and usually extended beyond R,,;, but nev-
er reaching R,,;; one midway between pter-
ostigma and apex of R,,;, triangular and at
most extended to R,,;; and a small trian-
gular or quadrate subapical spot often ex-
tended to vein R,,;. Cell r,,, usually with
2 small or 1 large marginal hyaline spots,
medial dark ray present (Fig. 1b), incom-
plete or absent (Fig. 1c—d). Basal cells hy-
aline. Cell cu, hyaline except apical margin
and sometimes a small subapical brown
spot touching Cu,. Legs: yellow. Male fore
femur no wider than that of female; in both
sexes with posteroventral row of 5 whitish
inflated setulae, the basal 2 relatively short-
er; also 2 dorsal rows of whitish inflated
setulae at most as long as 2 basal of pos-
teroventral row. Male fore tarsus with first
tarsomere as long as tarsomeres 2-4 togeth-
er, without unusual setation. Abdomen: In
both sexes microtrichia and setulae on ab-
dominal terga concolorous with those on
mesonotum. Female: tergite 2 with small
setulae on anterior margin, centrally bare
and with sparse setulae on posterior half;
setulae slightly increasing in density from
tergite 2 to 3 and uniformly dense on ter-
gites 4—6. Syntergosternite 7 (Fig. 2a)
black, 0.88—0.98 mm long, 0.45—0.53 mm
wide at base and 0.12—0.21 mm wide at
apex; with evenly distributed white setulae.
Eversible membrane 0.8—1.1 mm long,
scales as in Fig. 2b. Aculeus pale brown,
0.79—0.88 mm long, in lateral view straight
with tip slightly curved downward; tip tri-
angular, with 2 pairs of hairlike sensilla and
slightly notched at extreme apex (Fig. 2c).
One pair of spermathecae, ovoid, 0.49—0.62
mm long and 0.30—0.32 mm wide, surface
with papillae (Fig. 2d). Male: setulae on ter-
gites similar to female in size, density and
distribution. Sternite 5 with posterior mar-
gin concave (Fig. 2e). Epandrium with setu-
lae and microtrichia distributed as in Fig.
2f-g. Outer surstylus indistinguishably
fused to epandrium. Inner surstylus with 2
251
pairs of prensisetae; lateral prensiseta con-
ical, ca. % as long as mesa! prensiseta; be-
tween prensisetae a group of 2-3 setulae
and also a group of 3 setulae near base of
lateral prensiseta. Basiphallus 0.90—1.10
mm long; internal sclerites of distiphallus
0.20—0.23 mm long, apex of distiphallus
membranous, ca. % length of internal scle-
rites.
Egg: Ellipsoidal, shiny, white, 1.0 mm
long and 0.3 mm wide (n = 10). Chorion
with pattern of more or less hexagonal ar-
eas, more elongated on anterior end (Fig.
2h). Posterior pole tapered. Anterior pole
ends in nipple-shaped pedicel perforated
with several aeropyles (Fig. 21).
Larva: Mature larva (third instar) yellow,
globose, 2.65 + 0.1 mm long and 2.09 +
0.16 mm wide (+SD, n = 20) (Fig. 3a).
The most remarkable character is a poste-
rior area between posterior spiracles and
anal slit, which is more sclerotized and pale
brown. Sclerotized area on the ventral edge
with small indentation enclosing anal slit;
laterally some darker marks aligned in two
pairs of parallel lines at 30° angle with me-
dial line (Fig. 3b). Thoracic segments with-
out spinules or a few on third segment, ab-
dominal segments with conical spinules
more numerous on dorsum; spinules on the
dorsum of abdominal segments 7 and 8
more sclerotized, producing distinct dark
area. Surface of posterior sclerotized area
uniformly covered by conical spinules, rest
of caudal segment with spinules as on ab-
dominal segments. Cephalopharyngeal
skeleton as in Fig. 3c. Mandible short, with
two stout, apically rounded teeth. Labial
sclerite well developed. Hypopharyngeal
sclerite rectangular, elongate, ca. three
times as long as high. Parastomal bars fused
to both hypopharyngeal sclerite and tento-
ropharyngeal sclerite. Pigmented area of
dorsal cornu with posterior end bifid, ven-
tral cornu with ovoid window. Anterior spi-
racle with 3—4 papillae, felt chamber filled
with round reticulation (Fig. 3d). Posterior
spiracle located above medial horizontal
line of caudal segment (Fig. 3b). Dorsal spi-
252 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Trypanaresta valdesiana: A-D female (A) female terminalia; (B) scales of the eversible membrane;
(C), aculeus tip; (D) spermatheca; E-G male terminalia (E) sternite 5; (F) epandrium and surstyli, lateral view;
(G) epandrium and surstyli (proctiger not shown), posterior view ; (H) egg; (I) detail of pedicel. ac, aculeus;
em, eversible membrane; isu, inner surstylus; osu, outer surstylus; pro, proctiger; prs, prensisetae; s7, synter-
gosternite 7.
VOLUME 99, NUMBER 2
i)
Nn
i>)
Fig. 3. Trypanaresta valdesiana, third instar larva (A) habitus, lateral view; (B) posterior view; (C) cephal-
opharyngeal skeleton; (D) anterior spiracle; (E) posterior spiracle. sa, sclerotized area; an, annus; dc, dorsal
cornu; es, ecdysial scar; hs, hypopharyngeal sclerite; ih, intraspiracular hairs; Is, labial sclerite; md, mandibles:
pb, parstomal bar; re, reticulum; ri, rima; tr, trabeculae; vc, ventral cornu.
254
racular opening at right angle with medial
line, and at approximately 60° angle with
ventral spiracular opening (Fig. 3e). Upper
edge of rimae somewhat twisted, inner edge
with short and irregular trabeculae, never
forming crossbars; felt chamber with round
reticulation. Four groups of branched in-
traspiracular hairs, each branch blade end-
ing in acute tip, the 2 groups closer to cen-
tral rima with 1-3 branches, the other 2
groups with 4—6 branches (Fig. 3e).
Puparium: Puparium more cylindrical
than larva. Average length 2.8 + 0.27 mm,
maximum width 1.4 + 0.16 mm (n = 20).
The anterior extreme, including invagina-
tion scar, dark brown to black, the rest uni-
formly pale brown or rarely ivory. The
sclerotized area described for the larva
readily observed in puparium.
Etymology.—The epithet is an adjective
that refers to the Valdes peninsula, Chubut
province, Argentina, where the larvae were
originally collected.
Biology.—At the beginning of spring
(September), females presumably lay eggs
in immature heads of G. solbrigii. The
heads attacked by T. valdesiana are distin-
guished as they dry up prematurely and
never fully open (Fig. 4). The contents of
these “dry heads”’ are totally consumed and
the space is occupied by the larva. The ma-
ture larva is inside a cell with rigid walls
apparently formed by flower tissues ce-
mented by feces. No achenes are produced
in infested capitula. At the end of the
blooming season most larvae enter diapause
and remain inside the dried flower heads
until the next spring when they pupate and
adults finally emerge. In the population
studied near Puerto Piramide, about 5—10%
of the larvae pupariate near the end of the
blooming season (January), and behaves as
a bivoltine species. The adults that emerge
from those puparia are still able to find
some plants with young heads suitable for
Oviposition.
Natural enemies.—Two species of endo-
parasitic chalcidoids wasps, Epicatolaccus
strobeliae Blanchard (Pteromalidae) and
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. Flower head of Gutierrezia solbrigii in-
fested by Trypanaresta valdesiana.
Torymoides sulcius (Walker) (Torymidae),
were reared from larvae and puparia of T.
valdesiana. The former was known as a
parasitoid of the nonfrugivorous tephritid
Strobelia baccharidis Rondani (Blanchard
1940), and has been reared from immatures
of at least six species of florivorous tephri-
tids from Patagonia (Gandolfo, unpub-
lished). Torymoides sulcius was known as
a parasitoid of gall midges, and occurs from
Texas to South America (E.E. Grissell, pers.
comm.). In samples of flower heads of G.
solbrigii that were infested by larvae of T.
valdesiana and Trupanea patagonica (Bré-
thes), Torymoides sulcius was reared exclu-
sively from the former species. Both para-
sitoids killed, at the end of the 1994-95
growing season, 53% of the larvae and pu-
pae of T. valdesiana in a population near
Puerto Pirdamide (Gandolfo, unpublished).
Instead of leaving the bud through the apex,
as the flies do, the adult parasitoids make a
JESS)
VOLUME 99, NUMBER 2
'
'
‘
‘
’
'
‘
'
'
'
1
-~
]
a
x
oo ak
’
'
‘
'
'
1
‘
‘
ee es es ee
a alot y an nls ww ws A a ai Oy | aye!
1
'
Al
@ adults reared from flower
heads of Gutierrezia solbrigii
© adults reared from flower
heads of G. spathulata
References of Provinces
CH, Chubut
LP, La Pampa
ME, Mendoza
NE, Neuquén
RN, Rio Negro
Geographical distribution of Trypanaresta valdesiana.
Fig. 5.
256
circular hole on the upper half of the dry
bud.
Host plants.—Gutierrezia solbrigii Ca-
brera, G. mandonii (Schultz Bipontinus)
Solbrig and G. spathulata (Phil.) Kurtz.
Geographical distribution.—Argentina:
west to east Patagonia between 38° and 43°
S (Provinces of Chubut, Rio Negro, Neu-
quen, La Pampa and Mendoza) (Fig. 5).
ACKNOWLEDGMENTS
We thank L. De Santis (Museo de Cien-
cias Naturales, La Plata, Argentina) for the
identifications of the parasitoids and A. Ca-
brera (Instituto Darwinion, San Isidro, Ar-
gentina) for the identifications of Gutierre-
zia spp. We are also grateful to M. C. Her-
nandez for preparing the illustrations. We
thank E.E. Grissell, E C. Thomson, W.N.
Mathis, L.E. Carroll, G.J. Steck, and R.D.
Goeden for their reviews of the manuscript.
LITERATURE CITED
Blanchard, E. E. 1940. Descripci6n de dos himenop-
teros (Chalcid.) pardsitos de larvas de Strobelia
baccharidis Rond. Revista del Museo de La Plata
(Nueva serie) 2: 85—98.
Cabrera, A. L. 1971. Compositae, pp. 1-141. Jn Cor-
rea, M. N., ed., Flora Patagonica, Vol. 8, Part VII.
Col. Cientifica INTA, Buenos Aires, Argentina.
Cordo, H. A. and C. J. DeLoach. 1992. Occurrence
of snakeweeds (Gutierrezia:Compositae) and their
natural enemies in Argentina: Implications for bi-
ological control in the United States. Biological
Control 2: 143-158.
DeLoach, C. J. 1981. Prognosis for biological control
of weeds of southwestern U.S. rangelands, pp.
175-199. In Delfosse, E. S., ed., Proceedings of
the V International Symposium of Biological
Control of Weeds, Brisbane, Australia, 22—29 July
1980. CSIRO, Melbourne, Australia.
Foote, R. H. 1967. Family Tephritidae. Jn Vanzolini,
P. E. and N. Papavero, eds., A Catalog of the Dip-
tera of the Americas South of the United States.
Fasc. 57 Dept. Zool., Sec. Agr., Sao Paulo, Brazil.
Sil yoyo),
1980. Fruit fly genera South of the United
States (Diptera: Tephritidae). U. S. Depart. of
Agr., Technical Bulletin 1600, 79 pp.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Foote, R. H., E L. Blanc and A. L. Norrbom. 1993.
Handbook of the fruit flies (Diptera: Tephritidae)
of America North of Mexico. Cornell University
Press, Ithaca, NY. 571 pp.
Frias, D. 1988. Tephritis marisolae, nueva especie
chilena del género Tephritis Latreille (Diptera-Te-
phritidae). Revista Chilena de Entomologia 16:
77-81.
Hering, E. M. 1940. Neue Arten und Gattungen. Si-
runa Seva 1: 1-16.
Jenkins, J. and J. Turner. 1989. Revision of the Bac-
charis-infesting (Asteraceae) fruit flies of the ge-
nus Tephritis (Diptera: Tephritidae) in North
America. Annals of the Entomological Society of
America 82: 674—685.
Lane, M. A. 1985. Taxonomy of Gutierrezia (Com-
positae: Astereae) in North America. Systematic
Botany 10: 7—28.
McGinty, A. and T. G. Welch. 1987. Perennial broom-
weed and Texas ranching. Rangelands 9: 246—
249.
Norrbom, A. L. 1993. New species and the phyloge-
netic analysis of Euaresta Loew (Diptera: Tephrit-
idae), with a key to the species from the Americas
south of Mexico. Proceedings of the Entomolog-
ical Society of Washington 95: 195-209.
Norrbom, A. L. and K. C. Kim. 1988. Revision of the
schausi group of Anastrepha Schiner (Diptera: Te-
phritidae), with a discussion of the terminology of
the female terminalia in the Tephritoidea. Annals
of the Entomological Society of America 81: 164—
7/35
Solbrig, O. T. 1960. Cytotaxonomic and evolutionary
studies in the North American species of Gutier-
rezia (Compositae). Contributions of the Gray
Herbarium 188: 63 pp.
1966. The South American species of Gu-
tierrezia. Contributions of the Gray Herbarium
197: 3-42.
Steck, G. J. and R. A. Wharton. 1986. Descriptions
of immature stages of Eutreta (Diptera: Tephriti-
dae). Journal of the Kansas Entomological Society
59: 296-302.
Teskey, H. J. 1981. Morphology and terminology—
larvae, pp. 65-88. Jn McAlpine, J. F et al., eds.,
Manual of Neartic Diptera, vol. 1 Research
Branch, Agriculture Canada, Monograph 27, Ot-
tawa, Canada.
Torrel, L. A., K. Williams and K. C. McDaniel. 1990.
Economics of broom snakeweed control, pp. 113-—
139. In Huddleston E. W. and R. D. Pieper, eds.,
Snakeweed: Problems and Perspectives, Proceed-
ings, Symposium Nov. 9-10, 1989, Agricultural
Experimental Station Bulletin 751, New Mexico
State Univ., Las Cruces, New Mexico.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 257-266
REDESCRIPTION OF THE SUBGENUS HYGIA (EUCOLPURA) BREDDIN
(HEMIPTERA: COREIDAE: COLPURIND, WITH THE DESCRIPTION OF
TWO NEW SPECIES, AND A KEY TO THE KNOWN SPECIES
HARRY BRAILOVSKY AND ERNESTO BARRERA
Departamento de Zoologia, Instituto de Biologia UNAM, Apdo Postal No. 701-53,
México, 04510 D.F, México.
Abstract.—The subgenus Hygia (Eucolpura) Breddin is redescribed and two new spe-
cies H. (E.) heveli and H. (E.) melas from Borneo are described; H. (E.) speculatrix
(Breddin) is made a junior synonym of H. (E.) lugubris (Walker); habitus view illustrations
and drawings of the male and female genitalia are provided; a key to the known species
is included.
Key Words:
Borneo
The genus Hygia Uhler (1861) contains
ten subgenera, Australocolpura Brailovsky,
Caracolpura Breddin, Colpura Bergroth,
Eucolpura Breddin, Hygia Uhler, Micro-
colpura Breddin, Pterocolpura Blote,
Sphinctocolpura Breddin, Stenocolpura
Breddin and Trichocolpura Breddin, and
approximately 79 species, widely distrib-
uted in the Oriental Region throughout Ja-
pan, China, Taiwan, India, Burma, Assam,
Korea, Cambodia, Malacca, Sarawak (Bor-
neo), West Malaysia, Sumatra, Singapore,
Java, Philippines, Sulawesi, Mentawei,
New Guinea and Australia (Brailovsky
1993)
The present paper adds two new species
in Hygia (Eucolpura) whose members are
distinguished by having the genae laterally
armed, the pronotum nearly campanuli-
form, and the humeral angles angulately or
nearly prominent.
The following abbreviations are used for
the institutions cited in this paper: BMNH
(The Natural History Museum, London,
England); BPBM (Bernice P. Bishop Mu-
seum, Honolulu, Hawaii); DEI (Deutsches
Insecta, Heteroptera, Coreidae, Colpurini, Hygia (Eucolpura), new species,
Entomologisches Institut, Germany); IRNB
(Institut Royal des Sciences Naturelles,
Bruxelles, Belgique); NSMT (National Sci-
ence Museum, Tokyo, Japan); UNAM (Co-
lecci6n Entomolégica, Instituto de Biolo-
gia, Universidad Nacional Aut6énoma de
México); USNM (National Museum of Nat-
ural History, Smithsonian Institution,
Washington D.C.); ZIL (Zoological Insti-
tute, Leningrad); ZMUA (Zoologisches
Museum, Universiteit van Amsterdam,
Netherlands).
All measurements are given in millime-
ters.
Hygia (Eucolpura) Breddin
Colpura (Eucolpura) Breddin, 1900b: 202.
Hygia (Eucolpura) Blote, 1936: 35, 38.
Diagnosis.—The genus Hygia Uhler in-
cludes ten subgenera (Brailovsky, 1993),
two of which, H. (Colpura) Bergroth and
H. (Eucolpura) Breddin, have the genae
projecting into acute or obtuse teeth. In the
other subgenera, the genae are, simple with-
out teeth or lateral projections.
Hygia (Colpura) includes more robust
258
species, with the sharp teeth of genae clear-
ly projecting laterally and visible from
above; the pronotum is trapezoidal with the
anterolateral borders straight or nearly so,
and the humeral angles are very broadly
rounded. In AH. (Eucolpura), the teeth of
genae are obtuse, the pronotum is more or
less campanuliform, and the humeral angles
are angulate and prominent.
Redescription.—Head: Wider than long,
pentagonal, flat dorsally; tylus unarmed,
globose apically, extending anteriorly to ju-
gae, slightly raised in lateral view; jugae
unarmed, thickened, shorter than tylus; gen-
ae with obtuse teeth directed forward; an-
tenniferous tubercle unarmed; side of head
in front of eye unarmed, subparallel; anten-
nal segment I moderately robust, thickest,
slightly curved outward and longer than
head; segments II and III cylindrical, slen-
der; segment IV fusiform, slender; segment
II the longest, segment IV shortest and III
subequal to I; antennal segment IV longer
than length of head; ocelli well developed,
strongly elevated; preocellar pit deep; eyes
large, spherical, sessile; postocular tubercle
protuberant; buccula rounded, short, not
projecting beyond antenniferous tubercle,
with sharp spiny anterior projection; ros-
trum reaching anterior third of abdominal
sternite III or anterior third of V; mandib-
ular plate unarmed.
Thorax: Pronotum wider than long, cam-
panuliform, slightly declivent; collar wide;
frontal angles rounded or produced forward
as small lobe or conical teeth; humeral an-
gles rounded and either not or angulately
exposed, prominent and elevated; anterolat-
eral borders with anterior half convex and
posterior half oblique, straight; posterolat-
eral borders and posterior border nearly
straight; callar region slightly convex, well
developed with median longitudinal depres-
sion. Anterior lobe of metathoracic peritre-
me reniform, globose, posterior lobe sharp,
small.
Legs: Femora densely granulate, with
two rows of spines along ventral surface;
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tibiae with shallow sulcus, sometimes dif-
ficult to see.
Scutellum: Triangular, flat, longer than
wide; apex barely globose, subacute or
nearly flat and acute.
Hemelytra: Macropterous, reaching al-
most the apex of the last abdominal seg-
ment; claval suture evident; apical border
oblique, straight, with short apical angle,
not reaching middle one third of hemelytral
membrane.
Abdomen: Connexival segments higher
than abdominal terga: superior border of
connexiva serrate; posterior angle of each
connexival segment complete, except the
VI moderately exposed; abdominal sterna
with medial furrow extending to the ante-
rior margin of sternite V.
Integument: Body surface rather dull,
with mesosternum lateraly shining. Head
ventrally, pronotum, scutellum, clavus, co-
rium, thorax, and abdomen densely covered
with circular greyish-white farinose punc-
tures, with short decumbent silvery bristle-
like setae, intermixed with few long erect
hairs located on the abdominal sterna. Head
dorsally and callar region scarcely punctate
to smooth; male genital capsule and female
genital plates densely punctate, with long
erect or semierect setae. Pubescence of an-
tennae and femora short, mainly suberect,
on tibiae and tarsi longer and rather dense.
Male genitalia: Genital capsule: Postero-
ventral edge subtruncated, or shallowly
emarginated with lateral lobes short and
wider (Figs. 2, 6), or short and narrow (Fig.
4), or projected on a medium plate with lat-
eral lobes subacute (Fig. 3) or deeply emar-
ginated, with lateral lobes elongated (Fig.
5) or strongly bilobed with lateral lobes re-
markably wider (Fig. 1).
Female genitalia: Abdominal sternite
VII with plica and fissura; plica triangular,
wide, reaching anterior third (Fig. 10), or
almost medial third of sternite VII (Figs. 7—
9); gonocoxae I squarish, enlarged dorso-
ventrally, in lateral view with external face
entire, straight, and open ventrally; parater-
gite VIII quadrate, with spiracle visible;
VOLUME 99, NUMBER 2 259
Figs. 1-6. Hygia (Eucolpura), male genital capsule in caudal view. 1, H. (E.) melas. 2, H. (E). scrutatrix.
3, H. (E). lugubris. 4, H. (E). moesta. 5, H. (E). heveli. 6, H. (E). scrutatrix.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 7-10. Hygia (Eucolpura), female genital capsule in lateral view. 7, H. (E). melas. 8, H. (E). heveli. 9,
H. (E). scrutatrix. 10, H. (E). lugubris.
paratergite IX square, medium size, extend-
ing beyond the external face of gonocoxae
I and with the external margin of various
shapes (Figs. 7—10).
Type species.—Lybas lugubris Walker,
1871.
Historical notes.—Walker (1871) de-
scribed Lybas lugubris from three different
localities Singapore, Sarawak and New
Guinea and Lybas moestus from Java. The
third known species, Colpura speculatrix
(Breddin 1900a), was described from Bor-
neo and Sumatra. Breddin (1900b) revised
the Tribe Colpurini (= Pachycephalini, =
Lybantini), included in the genus Colpura
Bergroth (1894), the new subgenus Eucol-
pura, described the fourth species C. scru-
tatrix from Borneo, and transfered the last
three species into Eucolpura. Subsequently,
Breddin (1906) described C. (E.) dolens
from Sumatra and C. (E.) severa from Java
and synonymized C. (E.) speculatrix under
C. (E.) lugubris and H. (E.) severa under
H. (E.) moesta. Blote (1936) synonymized
the genus Colpura under Hygia Uhler
(1861), gave new records for H. (E.) lugub-
ris, H. (E.) moesta, H. (E.) scrutatrix, and
resurrected H. (E.) speculatrix to species
status.
Examination of type material for H. (E.)
lugubris (BMNH), AH. (E.) moesta
(BMNH), H. (£.) scrutatrix (DEI), and H.
(E.) speculatrix (DEI), study of undeter-
mined specimens, as well as the original de-
VOLUME 99, NUMBER 2
scription and drawings of the male genital
capsule of H. (E.) dolens, resulted in the
recognition of two new species from Bor-
neo and confirmation that H. (E.) specula-
trix is a junior synonym of H. (E.) lugubris.
On the general pattern of distribution for
the subgenus, we believe the New Guinea
locality for H. (E.) lugubris is erroneous.
Hygia (Eucolpura) melas Brailovsky and
Barrera, new species
@igs-.1)7 lsh)
Description.—Measurements: male first,
then female: Head length 2.08, 2.32; width
across eyes 2.26, 2.36; interocular space
12s, £20: mterocellar. space 0.52590/56;
preocular distance 1.36, 1.48; length anten-
nal segments: I, 3.36, 3.40; II, 4.28, 4.36;
Ill, 3.36, 3.48; IV, 2.36, 2.44. Pronotum:
Total length 3.32, 3.70; width across frontal
angles 1.80, 2.20; width across humeral an-
gles 5.08, 5.76. Scutellar length 2.28, 2.76;
width 2.12, 2.56. Total body length 15.36,
18.05.
Male.—Coloration: Head dark red, with
dorsal face of postocular tubercle, and an-
terior third of buccula yellow, with follow-
ing areas with bright orange reflections: ty-
lus, antenniferous tubercles, and posterior
third of buccula; antennal segments I to III
bright orange (base of I yellow) and IV pale
yellow with basal join and apex pale orange
brown; rostral segments I to III pale brown,
and IV pale orange yellow with apex pale
brown; pronotum pale brown red with cal-
lus, and humeral angles dark red to black;
scutellum, clavus, corium, connexival seg-
ments, thorax, and abdominal sterna pale
brown red with following areas yellow:
apex of scutellum, clearly discoidal spot lo-
cated on inner third of apical margin of co-
rium, posterior third of connexival seg-
ments III to VII, posterior third of pleural
sterna III to VII, and posterior border or
posterior margin of abdominal sterna V to
VII; anterior and posterior lobe of metatho-
racic peritreme creamy yellow; hemelytral
membrane dirty yellow, with veins and
large distal spot brown, with basal angle
261
black; dorsal abdominal segments I to III
pale orange yellow, and [V to VII dark red
with orange reflections; genital capsule dark
red; coxae brown red with pale orange re-
flections, and with apical third pale yellow;
trochanters pale yellow with small bright
orange spot located on inner third of apical
margin; femora and tibiae with three nar-
row longitudinal stripes dark brown red,
and three narrow stripes pale yellow; tarsi
bright orange yellow.
Male.—Structure: Rostrum reaching an-
terior to middle third of abdominal sternite
IV; frontal angles rounded; humeral angles
angulately exposed, and moderately prom-
inent. Genital capsule: Posteroventral bor-
der strongly bilobed, with lateral lobes re-
markably wider (Fig. 1).
Female.—Coloration: Similar to male.
Connexival segments VIII and IX, abdom-
inal segments VIII and IX, and genital
plates pale to dark brown red, with follow-
ing areas yellow: posterior third of connex-
ival segment VIII, posterior angle of para-
tergite VIII, and inner angle of gonocoxae
I. Genital plates. Parategite [IX square, me-
dium size, extending beyond external face
of gonocoxae I, and with external margin
entire and not folded (Fig. 7).
Variation.—l, rostral segment IV pale
orange; 2, hemelytral membrane yellow
with veins brown red, and basal angle
black; 3, dorsal abdominal segments I to V
or VI bright orange yellow, with punctures
darker; 4, scutellum and thorax dark red; 5,
femora and tibiae sometimes with dark
brown red and yellow stripes difficult to
segregate but always present; 6, posterior
lobe of metathoracic peritreme dirty yellow.
Type material.—Holotype: 6, Malaysia,
Borneo, Sandakan, Baker (without data)
(USNM). Paratypes: 2 6, 2 2, same data
as holotype (USNM, UNAM); | 2, N. Bor-
neo, Kuching, March 1900, Dyak (BMNH):;
1 2, Sarawak, Baram River, Gunong-Tam-
abo, 7 November 1920, J.C. Moulton
(UNAM); | 3, O. Borneo, Midden, 18 Feb-
ian ege5 eH Ge (Sicbers, (ZI); gle S
Borneo, M.O. Borneo Exp., Long Petak
262 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 11. Dorsal view of Hygia (Eucolpura) melas, male.
VOLUME 99, NUMBER 2
(450 mts.), 24 August to 9 September 1925,
H.C. Siebers (ZIL); 1 ¢, Sarawak, Ulu
Akar, November 1914, P. de EK (UNAM); 1
3, Sarawak, Matang, December 1898
(BMNH); 1 3d, 2 &, Sarawak, Mt. Matang,
21 December 1913, January to February
1914 (BMNH); 2 2, Malayan Peninsula,
Selangor (2000 mts.), C.B. Klon (without
data) (BMNH); 1 2, Malayan Peninsula,
Bukit Kutu, (3457 ft), 18 September 1930,
N.C.E. Miller (UNAM); 1 6, Malayan Pen-
insula, SE. Pahang Rompin Mining, 32—37
km., from Petoh, 24 January 1961, T.C.
Maa (BPBM); | ¢, Malayan Peninsula, Se-
langor, Gombak Valley, 20 October 1921,
H.M. Pendlebury (UNAM); | 6, Malayan
Peninsula, Pahang (without data) (IRNB).
Notes.—Hygia (E.) melas is easily distin-
guished by the strongly bilobed shape of
the posteroventral border of the male gen-
ital capsule, which is lateraly delimited by
two remarkably pronounced arms (Fig. 1)
and by having the external margin of the
paratergite [X entire and not folded (Fig. 7).
An additional condition is the peculiar col-
oration pattern of the femora, with three
dark brown, narrow, longitudinal stripes
and three pale yellow stripes.
Etymology.—From the Greek melas,
meaning black, and refering to the black
coloration of the humeral angles.
Distribution.—Known from the type lo-
calities.
Hygia (Eucolpura) heveli Brailovsky and
Barrera, new species
(Figs. 5, 8, 12)
Description.—Measurements: male first,
then female: Head length 2.24, 2.48; width
across eyes 2.32, 2.52; interocular space
LG; 12322 sinterocellar ‘space’ 052, ‘0:59:
preocular distance 1.38, 1.57; length anten-
nal segments: I, 3.92, 4.04; II, 5.04, 5.04;
Ill, 4.00, 3.88; IV, 2.72, 2.72. Pronotum:
Total length 3.04, 3.56; width across frontal
angles 1.72, 2.23; width across humeral an-
gles 4.52, 5.50. Scutellar length 2.28, 2.60;
width 1.76, 2.28. Total body length 15.69,
18.28.
263
Male.—Coloration: Head dark red with
dorsal face of postocular tubercle, and an-
terior third of buccula yellow, with follow-
ing areas with bright orange reflections:
apex of tylus, antenniferous tubercle, and
buccula; antennal segments I to III bright
orange, and IV pale yellow with basal join
bright orange; rostral segments I to III
bright chestnut orange, and IV bright or-
ange, with apical third darker; pronotum
with collar, anterior lobe, and humeral an-
gles black, and posterior lobe reddish
brown; scutellum dark reddish brown, with
apex pale yellow; clavus and corium pale
reddish brown with yellow discoidal spot
located on the inner third of apical margin
of corium; hemelytral membrane dirty yel-
low, with basal angle black, and apical third
with large brown blotch; connexival seg-
ments pale reddish brown, with posterior
third yellow; abdominal segments I to V
pale orange red, and VI to VII dark red;
thorax dark reddish brown, with acetabulae
bright chestnut orange; anterior and poste-
rior lobe of metathoracic peritreme creamy
yellow; abdominal sterna dark reddish
brown with following areas yellow: poste-
rior third of pleural sterna III to VII, and
posterior border of abdominal sterna V to
VII; genital capsule dark to pale reddish
brown, with lateral lobes bright orange;
coxae and trochanter bright yellow; femora
with two longitudinal stripes yellow and
two bright orange; tibiae and tarsi bright or-
ange.
Male.—Structure: Rostrum reaching
middle third of abdominal sternite IV or an-
terior third of V; frontal angles rounded; hu-
meral angles angulately exposed, and mod-
erately prominent. Genital capsule: Postero-
ventral border deeply emarginated, with lat-
eral lobes elongated (Fig. 5).
Female.—Coloration: Similar to male.
Connexival segments VIII and IX reddish
brown with posterior third yellow; abdom-
inal segments VIII and IX reddish brown;
genital plates reddish brown with internal
angle of gonocoxae I yellow. Genital plates.
Paratergite [IX square, conspicuously devel-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
264
(Eucolpura) heveli, male.
gia
Dorsal view of Hy
Fig. 12.
VOLUME 99, NUMBER 2
oped, extending beyond the external face of
gonocoxae I, and with external margin
slightly folded and curved (Fig. 8).
Variation.—The type material exhibits
some color variation: 1, postocular tubercle
entirely yellow; 2, space between eyes and
ocelli yellow; 3, hemelytral membrane pale
yellow with basal angle black; 4, femora
orange yellow with reddish brown granules.
Type material.—Holotype: d, Malaysia,
Sabah, 1 km., South of Kundasang, (1530
mts.), 28 August 1983, G.E Hevel and W.E.
Steiner (USNM). Paratypes: 2 2, same data
as holotype (UNAM, USNM); 2 &, B.N.
Borneo, Mt. Kinabalu, Kenokok (3300 ft),
23-25 April 1929 (BMNH); 2 2, B.N. Bor-
neo, near Kinabalu, Tenompok (4700 ft), 18
May 1929 (BMNH, UNAM); | 2, Borneo,
Midden. 16 October 1925, H.C. Siebers
(ZMUA).
Notes.—Hygia (E.) heveli, is similar to
H. (E.) lugubris (Walker) and to H. (E.) me-
las Brailovsky and Barrera, in having the
frontal angles of the pronotum rounded and
not produced forward as conical teeth, and
by the rostrum reaching anterior or middle
third of abdominal sternite [TV (occasionally
anterior third of V). Hygia (E.) heveli is rec-
ognized by the shape of the posteroventral
border of the male genital capsule (Fig. 5),
for having paratergite [IX conspicuously de-
veloped, with the external margin slightly
folded and curved (Fig. 8), and by the
length of the antennal segment II longer
than 5.00mm. The length of segment II in
other taxa is shorter than 4.50mm, and par-
atergite IX has the external margin con-
spicuously folded downward (H. (E.) lu-
gubris) (Fig. 10) or has the external margin
entire and not folded (H. (E.) melas) (Fig. 7).
Etymology.—Named for G. FE Hevel
(USNM).
Distribution.—Known from the type lo-
calities.
KEY TO THE KNOWN SPECIES OF HYGIA
EUCOLPURA
1. Hemelytral membrane with basal angle pale
vellows Drow eiete era ele ts moesta (Walker)
— Hemelytral membrane with basal angle black.
. Posteroventral border of the male cap-
sule simple, transversely straight with lateral
angles nearly rounded ........4a lolens (Breddin)
— Posteroventral border of the male genital cap-
sule not transversely straight, with rounded lat-
eralgangless(iiesaal—3)) (aeey te eee wie ene ee 3
3. Frontal angles of the pronotum produced for-
ward as small conical teeth. . .scrutatrix (Breddin)
— Frontal angles of the pronotum rounded. ..... 4
4. Posteroventral border of the male genital cap-
sule lateraly delimited by two remarkably pro-
nounced arms (Fig. 1); paratergite [IX with the
external margin entire and not folded (Fig. 7)
5 er Sep melas Brailovsky and Barrera new species
— Posteroventral border of male genital capsule
delimited by much shorter arms; paratergite IX
genital
i)
5. Posteroventral border of the male genital cap-
sule deeply emarginated, with lateral lobes
elongated (Fig. 5); paratergite [X with the ex-
ternal margin slightly folded and curved (Fig.
8) ... .heveli Brailovsky and Barrera new species
— Posteroventral border of the male genital cap-
sule with a short plate, having a sharp inwardly
curving spine on each side (Fig. 3); paratergite
IX with the external margin conspicuously
folded downward (Fig. 10) ... lugubris (Walker)
ACKNOWLEDGMENTS
The authors express their gratitude to the
following colleagues and institutions for the
loan of specimens and other assistance rel-
evant to this study: Janet Margerison-
Knight (BMNH); Gordon Nishida (BPBM),
Eckhard K. Groll (DEI); Jan van Stalle
(IRNB); Masaaki Tomokuni (NSMT); Tho-
mas J. Henry and G.FE Hevel (USNM); I.M.
Kerzhner (ZIL); and W. Hogenes (ZMUA).
Special thanks for the two anonymous re-
viewers.
LITERATURE CITED
Bergroth, E. 1894. Rhynchota Orientale. Revue d’ En-
tomologie 13: 152-164.
Bléte, H. C. 1936. Catalogue of the Coreidae in the
Rijksmuseum van Natuurlijke Historie, Part III.
Coreinae, Second part. Zoologische Mededeelin-
gen 19: 23-66.
Brailovsky, H. 1993. A revision of the tribe Colpurini
from Australia (Hemiptera-Heteroptera-Coreidae).
Memoirs of the Queensland Museum 34(1): 35—60.
Breddin, G. 1900a. Hemiptera gesammelt von Profes-
266 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
sor Kukenthal im Malayischen Archipel. Abhan- nales de la Societe Entomologique de Belgique
dlunger der Senckenbergischen Naturforschenden 50: 47-58.
Gesellschaft 25(1): 139-202. Uhler, P R. 1861. Rectification of the paper upon the
Breddin, G. 1900b. Materiae ad cognitionem subfam- Hemiptera of the North Pacific Expedition. Pro-
iliae Pachycephalini (Lybantini Olim). Ex Hem- ceedings of the Academy of Natural Sciences of
ipteris-Heteropteris, Fam. Coreidae. Revue d’ En- Philadelphia 1861: 286-287.
tomologie Caen 19: 194—217. Walker, E 1871. Catalogue of the specimens of He-
Breddin, G. 1906. Neue beitrage zur kenntnis von miptera Heteroptera in the collection of the British
Colpura Bergr., und verwandter Rhynchoten. An- Museum. Part IV, London, 211 pp.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 267-278
A NEW AGONOXENINE MOTH DAMAGING ARAUCARIA ARAUCANA
NEEDLES IN WESTERN ARGENTINA AND NOTES ON THE NEOTROPICAL
AGONOXENINE FAUNA (LEPIDOPTERA: GELECHIOIDEA: ELACHISTIDAE)
RONALD W. HODGES
Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart-
ment of Agriculture % National Museum of Natural History, MRC-168, Washington, DC
20560, U.S.A.
Abstract.—Characters of the adult, pupa, and larva of Araucarivora gentilii, new genus
and new species, are illustrated. The needle mining larvae cause conspicuous browning
to monkey-puzzle tree in west-central Argentina. The check list of Neotropical Lepidop-
tera is revised to accomodate genera and species newly assigned to Agonoxeninae. A list
of taxa removed from Agonoxeninae and placed in other families/subfamilies is provided.
Five genera are transferred to new family/subfamily associations. One new generic syn-
onym and 23 new combinations are proposed.
Key Words:
morphology, needle miner
In 1994 Mario Gentili (San Martin de los
Andes, Neuquén, Argentina) noticed severe
browning of needle tips of Araucaria ar-
aucana (Molina), K. Koch (Araucariaceae)
throughout the area. He enclosed several
twigs and waited for adults to emerge to
learn what was causing this damage. The
adults represent an undescribed species and
genus of the moth family Elachistidae, sub-
family Agonoxeninae. They are character-
ized as follows.
Araucarivora Hodges, new genus
Type species: Araucarivora gentilii
Hodges
Diagnosis.—Antenna without pecten;
forewing with well-developed stigma; valva
with free costal lobe, arms of gnathos free,
ventral surface of each with very fine spic-
ules; apophyses anteriores and apophyses
posteriores absent; larva lacking prolegs on
AS and A6.
Description.—Forewing (Fig. 2): broadly
Classification, check list, Cosmopterigidae, Chrysopeleiinae, life history,
lanceolate, stigma well developed; R;, Ry,
R, stalked, R; to anterior margin; M,—CuA,
separate; CuP present at margin; 1A and 2A
forked at base, 1A weak. Hindwing (Fig.
3): lanceolate; R, weak, to Sc at ¥% wing
length; Rs—M, separate; M, and CuA, con-
nate; CuA, from Y, wing length, 1A and 2A
forked at base, 1A weak; retinaculum of fe-
male diffuse, anteriorly directed scales be-
tween R and CuA. Abdomen: Sternum 2
with long, somewhat diffuse venulae, ven-
ulae extending anteriorly but not as apo-
demes (Fig. 6); segment 8 not modified.
Male genitalia (Figs. 4—5): Vinculum a
slender band, slightly expanded distally in
saccal region; aedeagus with recurved
flange at apex; juxta with pair of lateral
lobes, each setose at apex; gnathos expand-
ed distally; uncus narrowly triangular, an-
terolateral margin slightly excavated. Fe-
male genitalia (Fig. 7): Apophysis anterior-
is and posterioris absent; sternum 8 scler-
otized laterally and posterad of ostium
268
Fie: 1:
Araucarivora gentilii, habitus.
bursae; ostium bursae on anterior margin of
sternum 8; ductus seminalis arising just be-
yond base of ductus bursae; corpus bursae
finely spiculose; signum a lightly sclero-
tized, irregular plate with several inwardly
directed projections; papilla analis blunt,
densely setose on dorsal surface. Pupa
(Figs. 8, 24—27): antennae meeting at ¥,—%
their length, then diverging, exposing hind-
tarsi, extending to posterior margin of A4;
maxillary palpus, femur 1, femur 2 visible;
vertex, patagium, and anterior part of T2
coarsely spiculose; surface of wing finely
spiculose; lateral condyles present on ante-
rior margins of A5—7; well-developed “‘pu-
pal legs”’ present on A9, each with many
recurved setae at apex (Figs. 26-27); cre-
master absent, several recurved setae pres-
ent on AlO (Fig. 26). Larva (Figs. 9—23):
Frons extending %4 distance to epicranial
notch; stemmata well separated from each
other (Fig. 10); P, setae more distant from
each other than P, setae (Fig. 11); body
densely and coarsely spiculose; prothoracic
shield and pinacula heavily sclerotized, lat-
ter relatively large; secondary setae appar-
ently absent, except few on L group of A9;
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
spiracle on Tl approximately 2X diameter
of those on Al-—7, spiracle on A8 larger
than that on T1; tarsal setae setiform (Fig.
18); prolegs present on A3 and A4 (Fig.
20), absent on A5 and A6; crochets short,
in circle, 15—16 (Fig. 21); 12—15 crochets
on A1O0 in irregular line; Al, A2 with one
SV seta; A7—8 with one SV seta; A8 with
L1 dorsad of spiracle; A9 with 6—7 SV se-
tae; A9 with D2 setae on separate pinacula,
D1 seta slightly anterad of D2 and SD1.
Superficially, adults of Araucarivora
gentilii are similar to some species of Te-
tanocentria Rebel but differ in the lack of
a color pattern on the forewings. The pres-
ence of a stigma on the forewing, absence
of apophyses in the female, and absence of
prolegs on larval A5 and A6 are unique
character states in Agonoxeninae.
Araucarivora gentilii Hodges,
new species
Diagnosis.—The character states given
for the genus will serve to diagnose the spe-
cies.
Description.—Moth nearly uniformly
dark gray brown on dorsal surface (Fig. 1).
Head: Scales on haustellum pale yellow on
basal 4%; maxillary palpus gray on dorsal
surface, yellowish gray ventrally; labial pal-
pus slender, 2nd and 3rd segments approx-
imately equal in length, mainly white, first
segment with pale-gray scales on dorsal
surface, second segment with anterior mar-
gin broadly dark brown and a few dark-
brown scales laterally in mesial %, apex
dark brown, third segment dark brown an-
teriorly, white posteriorly; antenna dark
brown on dorsal surface at base, becoming
dark yellowish gray, ventral surface of
scape and basal Y, of shaft white/off-white.
Forewing: Nearly immaculate, dark gray
brown, tips of individual scales pale gray
on distal % of wing, slightly gray on basal
% of wing, base of each scale paler than
mesial part. Hindwing: Darker brown than
forewing, fringe nearly same color as wing;
prominent patch of white/pale-gray scales
extending from base along costal margin to
VOLUME 99, NUMBER 2 269
Stigma
Costal lobe of Valva
Tegumen
Vinculum
5
Figs. 2-6. Araucarivora gentilii. 2, Forewing [USNM slide 81596], line scale = 1.0 mm. 3, Hindwing
[USNM slide 81596], line scale = 1.0 mm. 4, Male genitalia, posteroventral view, aedeagus removed [USNM
81594], line scale = 0.5 mm. 5, Aedeagus [USNM 81601], line scale = 0.5 mm. 6, Abdominal sternum 2, d
[USNM slide 81601], line scale = 0.5 mm.
270 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Papillae Anales Frons
Maxilla
Antenna
Et
Ostium Bursae
Figs. 7-9. Araucarivora gentilii. 7, Female genitalia, ventral view [USNM 81595], line scale = 1.0 mm. 8,
Pupa, ventral view, line scale = 1.0 mm. 9, Mandible, mesial view, line scale = 0.4 mm.
slightly beyond % wing length and posterad Foreleg: Coxa and trochanter shining dark
into cell, narrowing at apex. Legs: Scales gray; femur mainly dark brown with pale
appressed to segments, except dorsal sur- gray on ventral margin; tibia and tarsus
face of hindtibia with long, slender scales. similarly colored, ventral margin nearly
VOLUME 99, NUMBER 2
Figs. 10-15. Araucarivora gentilii, larva. 10, Head capsule, lateroventral view, line scale = 100 pm. 11,
Head capsule, anterior view, line scale = 100 pm. 12, Head capsule, ventral mouth area, line scale = 100 pm.
13, Maxillary palpus, anteromesial view, line scale = 10 pm. 14, Antenna, anterior view, line scale = 10 pm.
15, Head and thorax, lateral view, line scale = 10 pm.
white. Mid- and hindlegs: Similar to foreleg gray with yellowish cast, anterior part of
but with more pale scales. Abdomen: Main- many segments with yellowish-brown
ly dark brown dorsally, scales on posterior scales. Wing length 5.8—6.0 mm.
margin of last two segments dark, slightly Types.—Holotype @. Argentina, Neu-
yellowish gray; ventral surface mainly pale quén, San Martin de los Andes; '7,, I 1995;
M. Gentili; ex needle mine Araucaria ar-
aucana. USNM [National Museum of Nat-
ural History, Washington]. Paratypes: 4 6,
11 2. Each with same data as for holotype;
USNM slides 81594-81596, 81600-81602.
Paratypes to The Natural History Museum,
London, BM(NH); M. Gentili; USNM. In
addition several larvae, pupae, and pupal
skins are preserved in alcohol in the
USNM.
Larvae make a large blotch mine in the
distal part of the very broad needles of Ar-
aucaria araucana. Frass remains in the
mine. Pupation occurs in the mine. Before
pupating the larva cuts a nearly round hole
through which the adult emerges. Because
less than % of each needle is eaten by the
larva, damage to the tree probably is mainly
cosmetic. However, because the needles re-
main on the tree for several years, the
heavily infested trees present a highly un-
attractive appearance.
TAXONOMIC SUMMARY OF NEOTROPICAL
MOTHS FORMERLY ASSIGNED TO
AGONOXENINAE
Agonoxeninae are relatively poorly
known, but I give a short summary on their
classification (Hodges, in press) in the
forthcoming volume on Lepidoptera in the
Handbook of Zoology. The major literature
is contained in Bottimer (1926), Bradley
(1966), Clarke (1965a, b), Common (1990),
Hodges (1978), Kasy (1976), Kuznetsov, N.
J. (1916), Riedl (1969), and Stehr (1987).
Worldwide 23 genera and 95 species are
known. Becker (1984b) listed nine genera
and 42 species for the Neotropical Region.
That document is modified to accomodate
transfer of genera among families and sub-
families and species among genera.
Amblytenes Meyrick, 1930 (type species
lunatica Meyrick) is here transferred from
Batrachedrinae.
Auxotricha Meyrick, 1931b (type species
ochrogypsa Meyrick) was transferred from
Depressariinae by Minet, 1990.
Glaucacna Forbes, 1931 (type species ir-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
idea Forbes) is here transferred from Ge-
lechiinae.
Nicanthes Meyrick, 1928 (type species
rhodoclea Meyrick) is known from a single
female. The genital characters suggest that
it belongs in Gelechioidea (possibly Gele-
chiinae) but not in Agonoxeninae.
Pammeces citraula Meyrick, 1922: 584
is here transferred to Homaledra (Batra-
chedrinae), NEW COMBINATION.
Pammeces crocoxysta Meyrick, 1922:
584 is here transferred to Homaledra (Ba-
trachedrinae), NEW COMBINATION.
Panclintis Meyrick, 1929 (type species
socia Meyrick) is here transferred to Gele-
chioidea (without family placement). Pan-
clintis socia has an ocellus; no Agonoxen-
inae have ocelli.
Prochola Meyrick, 1915 (type species
oppidana Meyrick) is here transferred to
Cosmopterigidae, Chrysopeleiinae. The
male genitalia of the lectotype of Prochola
oppidana (Clarke, 1965b: pl. 255) indicate
that the genus is a chrysopeleiine. It appears
to represent a valid genus. Examination of
the genitalia of one of the specimens in the
type series demonstrates that Meyrick had
a mixed series; the male genitalia of this
specimen (USNM genitalia slide 87690) in-
dicate that it is a species of Periploca
Braun. Eighteen species of Prochola of
Becker’s list (1984b: 43) are given new as-
signments in the following taxonomic sum-
mary.
Syntetrernis neocompsa Meyrick, 1933:
428 is here transferred to Scythris, Scythri-
didae, NEW COMBINATION.
Tocasta Busck, 1912 (type species pris-
cella Busck) was transferred from Coleo-
phorinae by Baldizzone, 1989.
Study of Eritarbes guttata Busck shows
that it belongs in Tetanocentria; it is trans-
ferred below. Eritarbes otiosa Walsingham,
1909, the type species of Eritarbes, is a
species of /thome; thus, Eritarbes is a ju-
nior synonym of /Jthome Chambers, 1875.
The result of this research is summarized
in the following revised check list of Neo-
VOLUME 99, NUMBER 2
» Ss
WF Loita
SP Se
US
Figs. 16-21. Araucarivora gentilii, larva. 16, Thoracic segment 2 with L setae, lateral view, line scale =
100 pm. 17, Thoracic segment 3, seta SD1, spinules, line scale = 10 pm. 18, Thoracic leg 2, apex, line scale
= 10 pm. 19, Abdominal segments 4—5, lateral view, lateral view, line scale = 100 pm. 20, Abdominal segments
4-5, ventral view, lateral view, line scale = 100 wm. 21, Proleg and crochets, abdominal segment 4, ventral
view, line scale = 10 pm.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 22-27. Araucarivora gentilii. 22, Larva, lateral view of abdominal segments 7—10, line scale = 100
uum. 23, Larva, dorsal view of abdominal segments 9—10, line scale = 100 pm. 24, Pupa, lateral view of segments
4-10, line scale = 100 pm. 25, Pupa, lateral view of segments 4—7, showing lateral condyles, line scale = 100
um. 26, Pupa, posteroventral view of segments 6-10, showing “‘pupal legs,” line scale = 100 pm. 27, Pupa,
apex of “‘pupal leg,” line scale = 10 pm.
VOLUME 99, NUMBER 2
tropical Agonoxeninae and pertinent Chry-
sopeleiinae.
AGONOXENINAE
Parametriotini
Araucarivora Hodges, new genus
gentilii Hodges, new species
Auxotricha Meyrick, 1931b: 189
ochrogypsa Meyrick, 1931b: 190 [lectotype
illustrated by Clarke, 1963: pl. 60]
Glaucacna Forbes, 1931: 369
iridea Forbes, 1931: 369 [illustrated by
Forbes, 1931]
Homoeoprepes Walsingham, 1909: 10
felisae Clarke, 1962: 375 [genital slides of
type series studied]
sympatrica Clarke, 1962: 381 [genital
slides of type series studied]
trochiloides Walsingham, 1909: 10 [illus-
trated by Clarke, 1962: Fig. 3; holotype
without abdomen; male unknown]
Microcolona Meyrick, 1897: 370
transennata Meyrick, 1922: 575 [lectotype
illustrated by Clarke, 1965b: pl. 251]
Nanodacna Clarke, 1964: 125
ancora Clarke, 1964: 126 [genital slides of
type series studied]
indiscriminata Clarke, 1965a: 93 [genital
slides of type series studied]
logistica (Meyrick, 1931c: 387, Colonopho-
ra)
vinacea (Meyrick, 1922: 574, Homaledra)
Pammeces Zeller, 1863: 152
albivitella Zeller, 1863: 152 [type not stud-
ied]
lithochroma Walsingham, 1897: 103 [type
not studied]
pallida Walsingham, 1897: 103 [type not
studied]
phlogophora Walsingham, 1909: 11 [type
not studied]
problema Walsingham, 1915: 458 [type not
studied]
Tetanocentria Rebel, 1902: 107
Aetia Chambers, 1880: 186, preoccupied by
Agassiz, 1847: 10
Platybathra Meyrick, 1912: 78
Parametriotes Kuznetsov, N. J. 1916: 628
Syntetrernis Meyrick, 1922: 573
Chaetocampa Busck, 1926: 804
agypsota (Meyrick, 1922: 580, Prochola),
NEW COMBINATION [type illustrated by
Clarke, 1965b: pl. 256]
catacentra (Meyrick, 1922: 582, Prochola),
NEw COMBINATION [lectotype illustrat-
ed by Clarke, 1965b: pl. 256]
catholica (Meyrick, 1917: 46, Prochola),
NEw COMBINATION [lectotype illustrat-
ed by Clarke, 1965b: pl. 256]
guttata (Busck, 1914: 1, Eritarbes), NEW
COMBINATION [genitalia of paratype
studied]
sollers (Meyrick, 1917: 46, Prochola), NEW
COMBINATION [lectotype illustrated by
Clarke, 1965b: pl. 259]
subtincta (Meyrick, 1922: 574, Syntetrer-
nis), NEW COMBINATION [lectotype il-
lustrated by Clarke, 1965b: pl. 259]
xiphodes (Meyrick, 1922: 574, Syntetrer-
nis), NEW COMBINATION [lectotype il-
lustrated by Clarke, 1965b: pl. 278]
Tocasta Busck, 1912: 4
Amblytenes Meyrick, 1930: 229, NEw
SYONYMY [Amblytenes is transferred
from Batrachedrinae]
lunatica (Meyrick, 1930: 230, Amblytenes),
NEw COMBINATION [genitalia of type
studied]
priscella Busck, 1912: 4 [genitalia of type
studied]
revecta (Meyrick, 1922: 582, Prochola),
NEW COMBINATION [lectotype illustrat-
ed by Clarke, 1965b: pl. 259]
Zaratha Walker, 1864: 789
[preliminary study of superficially similar
species from the Neotropics resulted in dis-
covery of 11 segregates based on genitalia. |
macrocera C. Felder & Rogenhofer, 1875:
pl. 140, Fig. 18 [Becker, 1984a: 248
276
illustrated a specimen from Costa Rica
that matches the abdomenless lecto-
type]
mesonyctia Meyrick, 1909: 17 [type not
studied]
pterodactylella Walker, 1864: 790 [lecto-
type illustrated by Becker, 1984a: 248]
niveiventris C. Felder & Rogenhofer,
1875: pl. 140, fig. 26 [type lost]
COSMOPTERIGIDAE
Chrysopeleiinae
Ithome Chambers, 1875: 93
Eritarbes Walsingham, 1909: 7, NEW SyYn-
ONYMY
fuscula (Forbes, 1931: 357, Prochola),
NEw COMBINATION [paratype exam-
ined]
otiosa (Walsingham, 1909: 7, Eritarbes),
NEw COMBINATION [paratypes exam-
ined]
Periploca Braun, 1919: 261
aedilis (Meyrick, 1915: 331, Prochola),
NEW COMBINATION [lectotype illustrat-
ed by Clarke, 1965b: pl. 255]
basichlora (Meyrick, 1922: 582, Prochola),
NEw COMBINATION [lectotype illustrat-
ed by Clarke, 1965b: pl. 255]
chloropis (Meyrick, 1922: 580, Prochola),
NEw COMBINATION [lectotype illustrat-
ed by Clarke, 1965b: pl. 256]
euclina (Meyrick, 1922: 583, Prochola),
NEw COMBINATION [lectotype illustrat-
ed by Clarke, 1965b: pl. 257]
obstructa (Meyrick, 1915: 332, Prochola),
NEW COMBINATION [holotype illustrat-
ed by Clarke, 1965b: pl. 257]
orphnopa (Meyrick, 1922: 582, Prochola),
NEw COMBINATION [holotype illustrat-
ed by Clarke, 1965b: pl. 258]
orthobasis (Meyrick, 1922: 582, Prochola),
NEW COMBINATION [lectotype illustrat-
ed by Clarke, 1965b: pl. 258]
prasophanes (Meyrick, 1922: 581, Procho-
la), NEW COMBINATION [holotype il-
lustrated by Clarke, 1965b: pl. 258]
semialbata (Meyrick, 1922: 581, Procho-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
la), NEW COMBINATION [lectotype il-
lustrated by Clarke, 1965b: pl. 259]
Prochola Meyrick, 1915: 331
oppidana Meyrick, 1915: 331 [lectotype il-
lustrated by Clarke, 1965b: pl. 255]
pervallata Meyrick, 1922: 581, MISPLACED
[lectotype illustrated by Clarke, 1965b:
pl. 258]
Siskiwitia Hodges, 1969: 10
holomorpha (Meyrick, 1931la: 282, Pro-
chola), NEW COMBINATION [holotype
illustrated by Clarke, 1965b: pl. 257]
Stilbosis Clemens, 1860: 170
Aeaea Chambers, 1874: 73.
Amaurogramma Braun, 1919: 261
ochromicta (Meyrick, 1922: 580, Procho-
la), NEW COMBINATION [lectotype il-
lustrated by Clarke, 1965b: pl. 257]
ACKNOWLEDGMENTS
I thank M. Gentili for discovering this
interesting moth and providing me with the
specimens that form the basis of this paper.
Special thanks go to D. Adamski who dis-
sected specimens, took the SEM photo-
graphs, and made the drawings. I thank D.
R. Davis (Department of Entomology,
Smithsonian Institution), S. Passoa (Animal
and Plant Health Inspection Service, PPQ),
A. S. Menke and D. R. Smith (Systematic
Entomology Laboratory) for reviewing the
manuscript.
LITERATURE CITED
Agassiz, L. [1847.] Nomenclatoris Zoologici, conti-
nens nomina systematica generum Animalium tam
viventium quam fossilium, secundum ordinen al-
phabeticum disposita, adjectis auctoribus, libris in
quibus reperiuntor, anno editionis, etymologia, et
familis, ad quas pertinent, in variis classibus. (In-
dex universalis). Fasc. 12. Jent and Gassmann, So-
loduri, viii + 393 pp.
Baldizzone, G. 1989. Contribuzioni alla conoscenza
del Coleophoridae. LVIII. Nuove sinonimie nel
genere Coleophora Hiibner (VII) (Lepidoptera).
Rivista Piemontese di Storia Naturale 10: 145—
154.
Becker, V. O. 1984a. The Neotropical Gelechioidea
VOLUME 99, NUMBER 2
(Lepidoptera) described by Cajetan Felder and
Alois E Rogenhofer. Systematic Entomology 9:
245-280.
1984b. Gelechioidea. Jn Heppner, J. B., ed.
Atlas of neotropical Lepidoptera. Checklist: Part
1 Micropterigoidea - Immoidea. W. Junk, The
Hague, xxvii + 112 pp.
Bottimer, L. J. 1926. Notes on some Lepidoptera from
eastern Texas. Journal of Agricultural Research
33: 797-819.
Bradley, J. D. 1966. A comparative study of the co-
conut flat moth (Agonoxena argaula Meyr.) and
its allies, including a new species (Lepidoptera,
Agonoxenidae). Bulletin of Entomological Re-
search 56: 453-472, figs. 1-38.
Braun, A. EF 1919. Notes on Cosmopterygidae [sic],
with descriptions of new genera and species (Mi-
crolepidoptera). Entomological News 30: 260—
264.
Busck, A. 1912. Descriptions of new genera and spe-
cies of microlepidoptera from Panama. Smithson-
ian Miscellaneous Collections 59(4): 1—10, pl. 1.
1914. New genera and species of microle-
pidoptera from Panama. Proceedings of the United
States National Museum 47 (2043): 1—67.
1926. Chaetocampa. In Bottimer, L. J., ed.,
Notes on some Lepidoptera from eastern Texas.
Journal of Agricultural Research 33: 804.
Chambers, V. T. 1874. Micro-Lepidoptera. Canadian
Entomologist 6: 72-77.
1875. Tineina from Texas. Canadian Ento-
mologist 7: 92-95.
. 1880. Descriptions of some new tineina, with
notes on a few old species. Journal of the Cincin-
nati Society of Natural History 2: 179-194.
Clarke, J. E G. 1962. Neotropical microlepidoptera, I
and II. Proceedings of the United States National
Museum 113: 373-388.
1963. Catalogue of the type specimens of
microlepidoptera in the British Museum (Natural
History) described by Edward Meyrick. vol. IV.
Trustees of the British Museum (Natural History),
London, 521 pp.
. 1964. A new genus and species from the Juan
Fernandez Islands. Proceedings of the Biological
Society of Washington 77: 125-126.
1965a. Microlepidoptera of Juan Fernandez
Island. Proceedings of the United States National
Museum 117: 1-106.
1965b. Catalogue of the type specimens of
microlepidoptera in the British Museum (Natural
History) described by Edward Meyrick. vol. V.
Trustees of the British Museum (Natural History),
London, 581 pp.
Clemens, B. 1860. Contributions to American lepi-
dopterology. No. 4. Proceedings of the Academy
of Natural Science Philadelphia 1860 [12]: 156—
174.
20
1990. Moths of Australia. Mel-
bourne University Press, Victoria, vi + 585 pp.
Common, I. E B.
Felder, C. and A. EF Rogenhofer. 1875. /n Felder, C.,
R. Felder, and A. E Rogenhofer, eds., 1264-1875.
Reise der Osterreichischen fregatte Novara um die
erde. Zoologischer Theil 29(2): 1—20, pl
Forbes, W. T. M. 1931. Supplementary report on the
heterocera or moths of Porto [sic] Rico. The Jour-
nal of the Department of Agriculture of Porto [sic]
Rico 4: 339-394.
Hodges, R. W. 1969. Nearctic Walshiidae notes and
new taxa (Lepidoptera: Gelechioidea). Smithson-
ian Contributions to Zoology 18: 1—30.
1978. Gelechioidea, Cosmopterigidae. The
Moths of America north Mexico 6.1: [iJ—x + |—
166, pl. 1-6.
. In press. Gelechioidea. /n Kristensen, N., ed.
Handbook of Zoology, Lepidoptera.
Kasy, E 1976. Uber die Familienzugehorigkeit einiger
‘Heliodinidae’—Gattungen (Lepidoptera). Anna-
len des Naturhistoriches Museums, Wien 80: 415—
430.
Kuznetsov, N. J. 1916. Description of Parametriotes
1—140.
theae, gen. n., sp. n. (Lepidoptera, Tineidae), a
enemy of the tea bush in Transcaucasia. Russkoe
Entomologicheskoe Obozrénie 15: 627—651. [In
Russian with extensive English summary. |
Meyrick, E. 1897. Descriptions of Australian micro-
lepidoptera. XVII. Elachistidae. Proceedings of
the Linnaean Society of New South Wales [22]:
297-435.
1909. Descriptions of micro-Lepidoptera
from Bolivia and Peru. Transactions of the Ento-
mological Society of London 1909: 13-43.
1912. Descriptions of South African micro-
lepidoptera. Annals of the Transvaal Museum 3:
63-83.
1915. Exotic Microlepidoptera 1(11): 321—
3),
. 1917. Exotic Microlepidoptera 2(2): 33-64.
1922. Exotic Microlepidoptera 2(18-19):
545-608.
1928. Exotic Microlepidoptera 3(13): 385—
416.
. [1929.]. The micro-Lepidoptera of the “St.
George’ expedition. Transactions of the Ento-
mological Society of London (December 31,
1928) 76: 489-521.
1930. Ergebnisse einer zoologischen Sam-
melreise nach Brasilien, inbesondere in das Ama-
zonasgebiet, ausgefiihrt von Dr. H. Zerny. V. Teil.
Micro-Lepidoptera. Annalen des Naturhistorisch-
en Museums, Wien 44: 223-268, pl. 1-2.
. 1931a. Reports of an expedition to Brazil and
Paraguay in 1926-27: microlepidoptera. Journal
of the Linnaean Society of London (Zoology) 37:
277-284.
278
1931b. Exotic Microlepidoptera 4(6): 161—
1927
1931c. Micro-lepidoptera from south Chile
and Argentina. Anales del Museo Nacional de
Historial Natural ‘“‘Bernardino Rivadavia’’, Bue-
nos Aires 36: 377-415.
1933. Exotic Microlepidoptera 4(14): 417—
448.
Minet, J. 1990. Remaniement partiel de la classifica-
tion des Gelechioidea, essentiellement en fonction
de caractéres pré-imaginaux (Lepidoptera Ditry-
sia). Alexanor 16(1989): 239-255.
Rebel, H. 1902. Lepidopteren aus Morea. Berliner En-
tomologische Zeitschrift 47: 83-110.
Riedl, T. 1969. Matériaux pour la connaissance des
Momphidae paléarctiques (Lepidoptera). Partie IX.
Revue des Momphidae européennes, y compris qu-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
elques espéces d’ Afrique du Nord et du Proche-Ori-
ent. Polskie Pismo Entomologiczne 39: 635-923.
Stehr, E W. 1987. Superfamily Gelechioidea. Jn Stehr,
E W,, ed. Immature Insects. Kendall/Hunt, Du-
buque, Iowa, 379-399.
Walker, E 1864. List of the specimens of Lepidopter-
ous insects in the collection of the British Muse-
um 29: 563-835.
Walsingham, L. [De Grey, T.]. 1897. Revision of the
West-Indian micro-Lepidoptera, with descriptions
of new species. Proceedings of the Zoological So-
ciety of London 1897: 54-183.
1909-1915. Biologia Centrali-Americana.
Insecta. Lepidoptera-Heterocera 4: xii + 1—482,
pl. 1-10.
Zeller, P C. 1863. Zwé6lf amerikanische nachtfalter.
Entomologische Zeitung zu Stettin 24: 136-155.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 279-289
LIMOMYZA, A NEW GENUS OF PRIMITIVE LIMOSININAE (DIPTERA:
SPHAEROCERIDAE), WITH FIVE NEW SPECIES FROM UN! TED STATES,
MEXICO, AND CENTRAL AMERICA
S. A. MARSHALL
Department of Environmental Biology, University of Guelph, Guelph, Ontario, Canada
N1G 2WI1.
Abstract.—The new genus Limomyza is described and included in the Limosininae
primarily on the basis of its wing venation, despite the overall close similarity of Limo-
myza species to members of the Copromyzinae. Five new species of Limomyza are de-
scribed and keyed, including the type species L. cavernicola from United States, L. venia,
L. archiptera and L. hirta from Mexico, and L. sharkeyi from Guatemala and Mexico.
Key Words:
The new genus Limomyza is diagnosed
on the basis of a complete cell cup, an open
cell bm, a mid ventral hind tibial bristle,
and a large inclinate orbital bristle between
the upper exclinate orbital and inner verti-
cal bristles. Limomyza challenges current
subfamilial concepts in the Sphaeroceridae,
because it presents an apparent mosaic of
characters previously used to diagnose the
subfamilies Copromyzinae and Limosini-
nae. Although Limomyza species are gen-
erally most similar to the Copromyzinae,
they show some of the reductions in wing
venation characteristic of the Limosininae.
The main lineages of Sphaeroceridae are
therefore briefly reviewed below prior to
description of Limomyza as a limosinine.
THE MAIN LINEAGES OF SPHAEROCERIDAE
The Sphaeroceridae can be broken into 3
lineages, as follows:
Tucminae.—This is the sister group to
the rest of the family, and contains only the
genus Tucma Mourgués-Schurter. Tucma
retains a well developed male tergite 6, the
loss of which appears to be a synapomor-
Sphaeroceridae, Diptera, taxonomy, Limosininae
phy of the rest of the family (Marshall
1996).
Limosininae.—The Limosininae (includ-
ing the vast majority of the species in the
Sphaeroceridae) is characterised by the loss
of crossvein bm-cu (i.e. cells bm and dm
fused; also the case in the putative copro-
myzines Palaeolimisina and Palaeocerop-
tera), reduction of the distal part of vein M
(also reduced in Palaeoceroptera), and usu-
ally the loss of cup, elongation of the arista,
and a head which is distinctly higher than
long.
Sphaerocerinae plus Copromyzinae.—
The Sphaerocerinae plus the Copromyzinae
form a clade characterised by an epandrium
with a deep (Copromyzinae) or complete
(Sphaerocerinae) lateral cleft. The epandrial
cleft is absent in one monophyletic group
currently included in the Copromyzinae (a
group including Lotophila Lioy, Borboril-
lus Duda, Dudaia Hedicke, Gymnometopi-
na Hedicke, Metaborborus Vanschu-
ytbroek, and Afroborborus Curran) but, as
interpreted by Norrbom and Kim (1984),
the epandrial cleft has been lost in these
genera. An epandrium that is completely di-
280
vided, presumably by development of the
epandrial cleft, is one of several apomor-
phic characters defining the Sphaerocerinae,
but the Copromyzinae is currently defined
only on plesiomorphic characters (cells bm
and cup complete, vein M reaching the
wing margin, costa reaching the tip of vein
M, long and telescoping female abdomen,
strongly setose tibia, simple, narrow-based
surstylus).
SUBFAMILIAL PLACEMENT OF LIMOMYZA
Limomyza species are very much like the
Copromyzinae in general habitus, with
long, telescoping female terminalia, incli-
nate inner orbital setulae, hind tibia with
dorsal and ventral bristles, richly setose tib-
iae, and a fully developed cell cup. The
male terminalia, with broadly fused, poste-
riorly lobate cerci; elongate, simple surstyli;
and a large epiphallus are also more similar
to copromyzines than most limosinines.
These, however, are all plesiomorphic char-
acters. One possible apomorphic character
linking Limomyza to the Copromyzinae is
the absence of a ring sclerite in the mem-
brane of the male sixth abdominal segment.
Tucma and most Limosininae have a trans-
lucent circular plate, usually ringed by a
darkened sclerite, in the membrane imme-
diately behind the sixth right spiracle of the
male abdomen. This ring sclerite (possibly
a large campaneiform sensillum) is com-
pletely absent from Limomyza, copromyzi-
nes, and sphaerocerines. Despite the super-
ficial similarity between Limomyza and
Compromyzines, the loss of the ring sclerite
is the only apparently apomorphic character
supporting a relationship between Limo-
myza and all or part of the sphaerocerine-
copromyzine lineage. On the other hand,
three characters seem to support the inclu-
sion of Limomyza in the Limosininae. The
head is almost 1.5 as high as long, in con-
trast to most Copromyzinae, in which the
head is usually almost as long as high.
There is considerable variation in head
shape within both groups, so the value of
this character is questionable. The other two
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
characters suggesting that Limomyza_ be-
longs in the Limosininae are venational
characters. It is largely on the basis of these
characters, the lack of a closed cell bm and
M not reaching the wing margin, that Li-
momyza is placed in the Limosininae.
Most other sphaerocerid genera with a
closed cell cup but without a closed cell bm
also belong in the Limosininae, and all such
genera were included in this group (as Lep-
tocerinae) by Hackman (1969). These in-
clude the Archileptocera genus-group (An-
omioptera Schiner, Archileptocera Duda,
and Palaeocoprina Duda), Hellerella Duda,
Palaeoceroptera Duda, Palaeolimosina
Duda, and Palaeoceroptera Duda. All of
these taxa except Anomioptera have been
treated as subgenera of Archileptocera, but
are elevated to genus and keyed by Mar-
shall (in press). Palaeolimosina and Palae-
oceroptera are like Limomyza in having a
copromyzine-like habitus. Palaeolimosina
is known only from a single, damaged fe-
male specimen with short antennae and M
reaching the wing margin, and probably be-
longs in the Copromyzinae. Palaeocerop-
tera is the genus most similar to Limomyza,
but the one species of Palaeoceroptera
known from males has an epandrial cleft,
which suggests that this genus also belongs
in the Copromyzinae, or at least in the
sphaerocerine-copromyzine clade. Both Pa-
laeoceroptera and Palaeolimosina are
known only from the southern hemisphere,
and lack the defining characters of Limo-
myza as listed below.
Following Cumming et al. (1995), and
Wheeler (1995), terminology used here for
some structures of the male and female ter-
minalia differs from that of McAlpine
(1981), and from this author’s earlier papers
on New World Sphaeroceridae. The term
gonostylus is used for the structure previ-
ously referred to as the paramere, and the
term subepandrial sclerite is used for the
sclerite previously called sternite 10. Mor-
phological terminology for female termin-
alia follows that of McAlpine (1981) with
the exception of the terminal tergites and
VOLUME 99, NUMBER 2
sternites, here called tergite and sternite 10
rather than epiproct and hypoproct.
Limomyza Marshall, new genus
Type species: Limomyza cavernicola,
New Species.
Defining and diagnostic characters.—Li-
momyza differs from all other sphaerocerids
with a cell cup but no basal medial cell in
having a mid-ventral hind tibial bristle, a
bristle between the orbital and inner vertical
bristles, and only one pair of (prescutellar)
dorsocentral bristles. The absence of setae
on the scutellar surface and the inclinate row
of inner orbital setulae are also diagnostic.
Generic description.—Body length usu-
ally 3-5 mm; body color black to dark red-
dish brown; most of frons and antennae red-
dish, vertex, orbits, interfrontal strips, ocel-
lar triangle, and middle of interfrontal area
silvery pruinose. One or 2 rows of inclinate
inner orbital setulae present midway be-
tween orbital and interfrontal bristles; a sin-
gle inclinate orbital bristle present above
upper exclinate orbital bristle, between up-
per orbital and inner vertical bristle (Fig. 5)
(other sphaerocerids have 0-2 exclinate or-
bital bristles only, although some authors
have called the inner vertical bristle an or-
bital bristle). Postocellar bristles cruciate;
ocellar bristles strong. Face deeply concave,
upper half carinate. Margin of labrum with
a flat, triangular, pruinose median portion;
clypeus black. First flagellomere apically
flattened but not tapered; arista dorsal, hairs
of medium length, arista slightly longer
than head height. Eye 0.9-—2.0 genal
height. Mid tibia setose, upper surface with
at least 3 anterodorsal bristles and 3 pos-
terodorsal bristles, and 2-3 distal dorsal
bristles (lower one large); lower surface
with a midventral bristle, apex ringed with
bristles. Hind tibia with a long, thin, distal
dorsal bristle, a short midventral bristle, and
a stout apical ventral bristle. Postpronotum
with 2 bristles. Dorsocentral bristles in a
single large prescutellar pair; acrostichal
setulae long, dense, in 8—15 rows between
dorsocentral areas. Scutellum broad, ca.
281
1.7X as broad as long, with 4 marginal bris-
tles, disc microtrichose only. Halter pale.
Wing with cell cup closed, vein Cu, strong;
crossveins r-m and dm-cu separated by less
than 1.5X length of dm; vein Cu, strong for
some distance beyond crossvein dm-cu;
R,,3; Sinuate to sharply bent; C ending at
apex of R,,;; alula narrow or of medium
width.
Male abdomen (Limomyza_ sharkeyi
known from @ only): Sternite 5 dark except
for a pale posteromedial area, densely se-
tulose near posteromedial area. Membrane
around and behind 6th right spiracle un-
modified. Synsternite 6+7 simple, laterally
setulose, dorsolaterally contiguous with
sternite 8; sternite 8 shining, mostly bare,
articulating with the shorter, setulose, epan-
drium at 2 widely separated points. Epan-
drium densely setose, with some long lat-
eral bristles. Subepandrial sclerite well-de-
veloped, simple. Cerci broadly fused to
form a very long subanal plate, ventrally
projecting to form lobes here referred to as
subcerci; posterolateral parts of epandrium
swollen behind surstylar base and variously
projecting ventrally as epandrial lobes; pos-
teroventral part of epandrial lobe overlap-
ping base of prominent subcercus. Surstylus
elongate, narrow base articulated with
epandrium and broad hypandrial arm. Hy-
pandrium forked posteromedially; hypan-
drial arms free from hypandrial apodeme.
Basiphallus with elongate epiphallus as
long as or almost as long as gonostylus; dis-
tal part of basiphallus very small. Gonos-
tylus narrow and parallel-sided distally,
apex blunt, slightly bent anteriorly; basal
part of gonostylus slightly broader than dis-
tal part, with a truncate anteroventral lobe.
Distiphallus with a dark, hooked basal part
and equal-length pairs of distal dorsal and
distal ventral lobes separated by a conspic-
uous concavity. Female genitalia (Limomy-
za hirta known from ¢ only): Tergites 1—5
broad, heavily sclerotized; tergites 6—8 very
narrow, pale, telescoped into preabdomen;
tergites 6 & 7 divided into entire anterior
parts and tripartite posterior parts; tergite 8
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-8.
1—4, Limomyza archiptera, male. 1, Terminalia, posterior. 2, Terminalia, left lateral, hypandrium
removed. 3, Phallus and associated structures. 4, Sternites 5—7. 5, L. cavernicola, head. 6, L. archiptera, wing.
7, L. cavernicola, wing. 8, L. venia, wing. Abbreviations: basi = basiphallus; disti = distiphallus; e.1. = epandrial
lobe; gon = gonostylus; S5 = sternite 5; sc = subcercus; sur = surstylus; S6+7 = synsternite 6+7; cs2 =
second costal sector; cs3 = third costal sector; vti = inner vertical bristle; ior = inclinate orbital bristle; eor =
exclinate orbital bristles.
entirely tripartite, lateral parts shining and
relatively dark. Tergite 10 small, with 2
small bristles; cerci setulose and apically
setose. Sternite 8 bipartite, each half pos-
teriorly setulose and with 2-3 bristles; ster-
nite 10 uniformly, sparsely setulose, with 2
longer bristles. Spermathecae (3) oval to
elongate, with transverse wrinkles, strongly
tapered and usually curved basally.
Etymology: The name Limomyza is from
a combination of the names of the type gen-
era of Limosininae and Copromyzinae, and
alludes to the copromyzine habitus of this
limosinine genus.
VOLUME 99, NUMBER 2
BIOLOGY AND DISTRIBUTION OF LIMOMYZA
Collection records suggest that Limomy-
za species are primarily coprophagous, and
that most species in the genus are associ-
ated with high elevations (over 2000 m) in
Mexico and Central America. Limomyza
cavernicola, which occurs in eastern and
central United States, has been collected in
caves.
KEY TO SPECIES OF LIMOMYZA
1. Eye very small, height less than genal height.
Back of head densely setose, postocular bris-
tles in double row on lower half and triple row
on upper half. Proximal half of mid tibia with
anterior and anterodorsal rows of bristles. Sur-
stylus without row of bristles at apex (Figs. 16,
18). Known only from type locality in Duran-
TOMMEXICON | oe. J Limomyza hirta, new species
— Eye larger, height at least 1.5 genal height.
Postocular bristles forming a single row below
and a double row near top of eye. Mid tibia
with only anterodorsal and posterodorsal bris-
tles proximally. Surstylus of known males with
an apical row of stout bristles (Figs. 1, 10, 22).
USA, Mexico and Central America. ........ 2
2. Body length ca. 4 mm. Inclinate interfrontal
setulae between orbital and interfrontal bristles
extending at least up to level of upper inter-
frontal. Four large posterodorsal bristles on
mid tibia. Pruinosity along posterior part of
gena divided into small dorsal and extensive
ventral parts. Mexico (Chiapas) and Guatemala
(Sronly) ie Limomyza sharkeyi, new species
— Body length less than 3.5 mm. One row of 2—
4 inclinate interfrontal setulae between lower
orbital and interfrontal bristles; no setulae on
upper half of frons. Three large posterodorsal
bristles on mid tibia. Pruinosity continuous
alons poOstenlon partoOL Sena... see oe 3}
3. Surstylus slender, width at 4 subequal to width
at *%4; distal comb on posterior surface and
twice as long as maximum surstylar width (Fig.
21). Epandrial lobe acute. Mexico
5 i Cece aera ns Limomyza venia, new species
— Surstylus slightly expanded and flattened dis-
tally, distal comb on posteroventral or ventral
surface and less than twice as long as maxi-
mum surstylar width. Epandrial lobe blunt,
strongly overlapping subcercus. ........... 4
4. Second costal sector greater than 1.5 third
costal sector; R2+3 weakly sinuate (Fig. 7).
Distal comb of surstylus slightly longer than
maximum surstylar width (Fig. 9). Eye ca.
1.6X genal height. USA
Si eee Limomyza cavernicola, new species
283
— Second costal sector less than 1.3 third costal
sector; R2+3 strongly sinuate (Fig. 6). Distal
comb of surstylus subequal to maximum sur-
stylar width (Fig. 2). Eye ca. 2.1>
height Mexicomearrse a.) ie. -
genal
Limomyza archiptera Marshall,
new species
(Figs. 1—4, 6)
Body length ca. 3.0 mm; body Color
black to dark reddish brown; tarsi, apices of
tibiae and halters pale. Interfrontal bristles
in 4 long, equal pairs; a row of 2-3 incli-
nate inner orbital setulae present midway
between orbital and interfrontal areas, up-
permost setula at level of lower orbital bris-
tle. Face deeply concave, upper half of face
silvery pruinose. Eye 2.1 genal height;
gena shining anterodorsally, silvery prui-
nose posteriorly and ventrally, pruinose
area with a large anterior bristle and several
setulae. Katepisternum with two large dor-
sal bristles, posterior bristle twice as long
as anterior. Acrostichal bristles long, in 8—
10 rows between dorsocentral areas. Wing
with R2+3 conspicuously but gradually
bent at basal third; second costal sector
1.2X length of third; distance between
crossveins r-m and dm-cu 1.4 length of
dm-cu; alula narrow.
Male abdomen: Epandrium densely se-
tose, with long dorsolateral and posteroven-
tral bristles. Subanal plate broad, posterior
part with conspicuous, setose medial ven-
tral lobes (subcerci) with one bristle prox-
imally, distally with 3 posteromedial bris-
tles and 4—5 smaller medial setae; epandrial
lobes overlapping subcerci basally and dis-
tally. Surstylus elongate, narrow base artic-
ulated with epandrium and broad hypandri-
al arm; anteriorly curved at %, distal part
dark with dense row of 6—8 stout, dark bris-
tles, row shorter than maximum surstylar
width.
Holotype (¢, CNC) and 13 Paratypes (4
3,9 2, GUE): MEXICO. GUERRERO, 4
mi W Mazatlan, 4800’, 30.viii—5.ix.1971,
284 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 9-15. Limomyza cavernicola. 9, Male terminalia, left lateral. 10, Male terminalia, posterior. 11, sternites
5—7, male. 12, Phallus and associated structures. 13, Female abdomen, ventral. 14, spermathecae. 15, Female
abdomen, dorsal.
VOLUME 99, NUMBER 2
Figs. 16-21.
hypandrium removed. 19, Sternites 5—7. 20, Phallus and associated structures. 21, L. sharkeyi, spermathecae.
Oak, tropical deciduous forest, human
dung, A. Newton.
Other paratypes: MEXICO. MEXICO. 1
mi E Ixtapan de la Sal, 6200’, km 78,
31.viii—6.ix.1971, tropical deciduous forest,
dung trap, A. Newton (1 6, 1 2, GUE).
Etymology: The name archiptera refers
to the primitive wing venation of the genus
as a whole.
Comments: Limomyza archiptera is ex-
ternally similar to Limomyza venia, differ-
ing primarily in features of the male geni-
talia. The surstyli of these species are dis-
tinctly different.
16-20, Limomyza hirta, male. 16, Terminalia, posterior. 17, Hind leg, left. 18, Terminalia, left,
Limomyza cavernicola Marshall,
new species
(Figs. 5, 7, 9-12)
Body length ca. 3.0 mm; body Color
black to dark reddish brown; tarsi, apices of
tibiae and halters pale. Interfrontal bristles
in 4—5 long, equal pairs; a row of 3—4 in-
clinate inner orbital setulae present midway
between orbital and interfrontal strips, up-
permost setula at level of lower orbital bris-
tle. Face deeply concave, upper half silvery
pruinose. Eye 1.6X genal height; gena shin-
ing anterodorsally, silvery pruinose anter-
oventrally, pruinose area with a large an-
286 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 22-28. Limomyza venia. 22, Male terminalia, posterior. 23, Male terminalia, left lateral. 24, Sternites
5-7, male. 25, Phallus and associated structures. 26, Female abdomen, ventral. 27, Spermathecae. 28, Female
abdomen, dorsal.
VOLUME 99, NUMBER 2
terior bristle and several setulae. Katepis-
ternum with two large dorsal bristles, pos-
terior bristle twice as long as anterior.
Acrostichal setulae in about 8—10 rows be-
tween dorsocentral areas. Wing with R2+3
sinuate, gradually curved up to costa near
apex, second costal sector 1.7 length of
third; distance between r-m and dm-cu
1.6X length of dm-cu; alula narrow.
Male abdomen: Epandrium densely se-
tose, with long dorsolateral and posteroven-
tral bristles. Subanal plate broad, posterior
part with conspicuous, setose ventral lobes
(subcerci) medially, epandrial lobes ven-
trally rounded, pale, basally overlapping
subcercus; subcercus with | posterior bris-
tle basally, distally with 3 posterodorsal
bristles and a row of ca. 9 medial setae.
Surstylus gradually curved anteriorly, distal
part broad, with inner row of 13-14 stout,
dark bristles, row ca 1.5 times as long as
maximum surstylar width.
Holotype (3, CNC): UNITED STATES.
KENTUCKY. Edmonson Co., Mammoth
Cave National Park, Running Br., 17-—
25.vi.1973, forest dung trap, S. Peck.
Paratypes: UNITED STATES. COLO-
RADO. Larimer Co., 5,8,22.viii,30.i1x.1996,
S. Fitzgerald (8 6,4 2, CSU). MISSOURI.
Texas Co., 10.5 mi NW Licking, unnamed
cave, 14.vi.1989, J.E. Gardner (3 6, 3 @,
GUE); OKLAHOMA. Caddo Co., 0.5 mi.
S. Hinton, Redrock Canyon State Park, 2—
3.vili.1984, dung trap, B.V. Brown (1 6,
GUE); Murray Co., Chickasaw National
Recreation Area, 24.v.1991, J.E. Swann (1
3d, GUE).
Etymology: The specific name refers to
the association of L. cavernicola with
caves.
Comments: Limomyza cavernicola is ex-
ternally very similar to Limomyza venia and
Limomyza archiptera. These species can be
most reliably separated on the basis of the
surstylus, but L. cavernicola also differs
from L. venia and L. archiptera in having
relatively small eyes, presumably associat-
ed with its hypogean habits. Limomyza hir-
ta, a species of unknown biology which is
287
easily distinguished from L. cavernicola by
characters given in the key, also has very
small eyes.
Limomyza hirta Marshall, new species
(Figs. 16—20)
Body length ca. 5.0 mm; body Color
black to dark reddish brown; antennae and
most of pleuron reddish. Interfrontal bris-
tles in 6-7 long, equal pairs; 2 rows of in-
clinate inner orbital setulae present midway
between lower orbital and interfrontal bris-
tles, inner row of ca 8 bristles extending up
to level of ocellar bristle. Postocellar bris-
tles convergent but equally long divergent
postocular bristles present immediately be-
hind postocellar bristles (the postocular
bristles form a dense double or triple row
behind the eye then extend across the back
of the head as a sparse single row). Face
deeply concave, pruinose; lateral margin of
face and clypeus black. Eye 0.9X genal
height; gena entirely pruinose, with a large
anterior bristle and several setulae forming
4 rows on ventral half. Katepisternum with
a large posterodorsal bristle and 2 very
small anterodorsal bristles less than % as
long as posterodorsal bristle. Upper surface
of mid tibia with a row of anterior bristles
paired with the usual anterodorsal bristles
on the proximal half of the tibia. Acrosti-
chal setulae long, in about 15 rows between
dorsocentral areas. Wing with R2+3 con-
spicuously but gradually bent at basal third;
second costal sector 2.2 length of third
costal sector; distance between r-m and
dm-cu 2.6 length of dm-cu.
Male abdomen: Epandrium densely se-
tose, with long dorsolateral bristles. Suban-
al plate very broad, posterior part with a
prominent, microsetulose median process;
subepandrial sclerite with a smaller, bare,
median process. Posterolateral corners of
epandrium forming subquadrate, setose
lobes (epandrial lobes): narrow posteroven-
tral lobes (subcerci) overlapped by epandri-
al lobes, bent posteriorly, parallel sided,
distally rounded with ca 8 bristles. Sursty-
lus simple, elongate-triangular, with long
288
anterior bristles. Gonostylus narrow and
gradually tapering distally, apex blunt, bi-
carinate.
Holotype (6, CNC): MEXICO. DU-
RANGO. 10 mi W El Salto, 13.vii.1964,
H.F Howden.
Etymology: The specific name refers to
the hirsute appearance of this large, distinc-
tive species.
Comments: Limomyza hirta can be easily
separated from congeners by the double
row of postocular bristles, the extra row of
anterior mid tibial bristles, the large size,
the distinctive wing venation with a very
long second costal sector, or by the sursty-
lus which lacks the apical comb row char-
acteristic of congeners.
Limomyza sharkeyi Marshall,
new species
(Fig. 21)
Body length ca. 4.0 mm; body Color
black to dark reddish brown; face and lower
frons reddish, antennae and tarsomeres
pale, orange; halter very pale. Interfrontal
bristles in 4 long, equal pairs; 2 rows of
inclinate inner orbital setulae present mid-
way between lower orbital and interfrontal
bristles, inner row of ca 6 bristles extending
up to level of ocellar bristle. Postocellar
bristles convergent. Postocular bristles in a
dense single row behind the eye and a dou-
ble row above the eye. Face slightly con-
cave, pruinose on upper half; lateral margin
of face and clypeus brown. Eye 2.0 genal
height; gena silvery pruinose on lower half,
upper half shining except for small postero-
dorsal pruinose area, with a large anterior
bristle and several setulae forming 4 rows
on ventral half. Katepisternum with a large
posterodorsal bristle and an anterodorsal
bristle % as long as posterodorsal bristle.
Upper surface of mid tibia with small prox-
imal anterodorsal and posterodorsal bristles
in addition to the usual 5 anterodorsal and
4 posterodorsal bristles, and one small an-
terior bristle at middle. Acrostichal setulae
long, in about 10 rows between dorsocen-
tral areas. Wing with R2+3 conspicuously
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
but gradually bent in basal third; second
costal sector 1.5 length of third; distance
between crossveins r-m and dm-cu twice as
long as length of dm-cu; alula of medium
width. Spermathecae elongate, with long
straight bases in contrast with the short,
curved bases of known congeneric females.
Holotype (2, CNC): GUATEMALA.
SAN MARCOS. San Antonio, 8000’, Sa-
catepequez, 29.1x.1987, M. Sharkey.
Paratype: MEXICO. CHIAPAS, Muni-
cipio El Porvenir, between El Porvenir and
Siltepec, N. Slope Cerro Male, 2134-743
m, 19-1x.1976, D.E. and J.A. Breedlove (1
PCAS):
Etymology: This species is named after
Mike Sharkey, who has generously allowed
me to study the flies from his insect trap
catches.
Comments: Although it is generally un-
desirable to describe species of Sphaerocer-
idae from females only, these large Limo-
myza specimens are obviously different
from all congeners. The chaetotaxy of the
mid tibia, size, spermathecae, pruinosity of
the gena, and inner orbital bristles all ap-
pear to be diagnostic, although the sper-
mathecae are unknown for one congener (L.
hirta).
Limomyza venia Marshall, new species
(Figs. 8, 22—28)
Body length ca. 3.0 mm; body color
black to dark reddish brown; tarsi, apices of
tibiae and halters pale. Interfrontal bristles
in 4—5 long, equal pairs; a row of 3—4 in-
clinate inner orbital setulae present midway
between orbital and interfrontal areas; up-
permost setula at level of lower orbital bris-
tle. Face deeply concave, upper half of face
silvery pollinose. Eye 2.0 genal height;
gena shining anterodorsally, silvery prui-
nose anteroventrally, pruinose area with a
large anterior bristle and several setulae.
Katepisternum with two large dorsal bris-
tles, posterior bristle twice as long as an-
terior. Acrostichal setulae in about 8—10
rows between dorsocentral areas. Wing
with R2+3 sharply bent up at basal third;
VOLUME 99, NUMBER 2
second costal sector 1.3—1.4x< as long as
third; distance between crossviens r-m and
dm-cu 1.2—1.3X length of dm-cu; alula nar-
row.
Male abdomen: Epandrium densely se-
tose, with long dorsolateral and posteroven-
tral bristles. Subanal plate broad, posterior
part with conspicuous, setose ventral lobes
(subcerci) medially, epandrial lobes ven-
trally narrow, acute, pale; subcercus with |
posterior bristle basally, distally with 3 pos-
terodorsal bristles and a row of ca. 9 medial
setae; epandrial lobe not conspicuously
overlapping subcercus basally. Surstylus
elongate, sharply bent at %, distal part with
inner row of 13—14 stout, dark bristles, row
over twice as long as maximum surstylar
width.
Holotype (6, CNC) and 90 Paratypes (48
6. 42 2. GUE;, FLD); MEXICO: MEXI-
CO. 1 mi NE Tenancingo, 7100’, 31.viii—
6.1x.1971, Oak -Pine, human dung trap, A.
Newton.
Other paratypes: MEXICO. MORELOS.
4 mi W Tres Cumbres, 9000’, 29.viii—
4.1x.1971, Oak, human dung trap, A. New-
tons(G.c. 3 2; GUE):
Etymology: The name of this species is
to be considered an arbitrary combination
of letters.
ACKNOWLEDGMENTS
Although most of the specimens exam-
ined are in the University of Guelph Col-
lection (GUE), the loan of material from the
following institutions is acknowledged: Ca-
nadian National Collection, Ottawa (CNC);
Colorado State University, Ft. Collins
(CSU); Field Museum, Chicago (FLD);
289
California Academy of Sciences, San Fran-
cisco (CAS), and the Museum of Zoology,
University of Lund, Sweden (LUND). I
thank Dr. Allen Norrbom for stimulating
discussions about character state distribu-
tion within the basal lineages of the Sphaer-
oceridae. Many thanks to Drs. B. Sinclair,
J. Rohacek, and T. Wheeler for reviewing
the manuscript, and to Rebecca Langstaff
for preparing the illustrations.
LITERATURE CITED
Cumming, J. M., B. J. Sinclair, and D. M. Wood.
1995. Homology and phylogenetic implications
of male genitalia in Diptera-Eremoneura. Ento-
mologica Scandinavica 26: 120-151.
Hackman, W. 1969. A review of the zoogeography
and classification of the Sphaeroceridae (Borbor-
idae, Diptera). Notulae Entomologicae XLIX,
1969: 193-210.
McAlpine, J. F 1981. Morphology and terminology—
Adults, pp. 9-63. In McAlpine, J. EF et al, eds.
Manual of Nearctic Diptera, Agriculture Canada
Monograph 27, 674 pp.
Marshall, S. A. 1996. Tucma fritzi, a new species in
the enigmatic genus Tucma Mourgués-Schurter
(Diptera; Sphaeroceridae; Tucminae, new subfam-
ily). Studia Dipterologica 3: 283-288.
Marshall, S. A. In press. A revision of Archileptocera
and related genera, including Anomioptera Schi-
ner, Palaeocoprina Duda, and Archileptocera
Duda, with a key to sphaerocerid genera with sim-
ilar wing venation and a description of a new spe-
cies of Palaeoceroptera Duda (Diptera, Sphaero-
ceridae). Journal of Natural History.
Norrbom, A. L. and K. C. Kim. 1984. The taxonomic
status of Lotophila Lioy, with a review of L. atra
(Meigen) (Diptera, Sphaeroceridae). Proceedings
of the Entomological Society of Washington 86:
305-311.
Wheeler, T. A. 1995. Systematics of the New World
Rachispoda Lioy (Diptera: Sphaeroceridae): mor-
phology, key to species groups, and revisions of
the atra, fuscipennis, limosa and vespertina spe-
cies groups. Journal of Natural History 29: 159—
230.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 290-293
A NEW SPECIES OF NEARCTIC PERLESTA (PLECOPTERA: PERLIDAE)
FROM VIRGINIA
RALPH FE KIRCHNER! AND Boris C. KONDRATIEFF
(RFK) Department of the Army, Huntington District Corps of Engineers, Water Quality
Section, 502 8th Street, Huntington, WV 25701, U.S.A.; (BCK) Department of Ento-
mology, Colorado State University, Fort Collins, CO 80523, U.S.A.
Abstract.—A new species of stonefly from southwestern Virginia is added to the Ne-
arctic Perlesta, which presently includes 14 species. Perlesta teaysia, new species, is
described from the adult male, adult female, egg, and mature nymph. Diagnostic features
are supported by illustrations and SEM photomicrographs.
Key Words:
Stark (1989) revised and keyed 12 spe-
cies in the Perlesta placida (Hagen) com-
plex. Poulton and Stewart (1991) and Stark
and Rhodes (1997) have added two addi-
tional species to the Nearctic list.
A distinctive new species of Perlesta was
reared by the senior author in 1979. How-
ever, not until recently did the adult male
become available with the extruded penis
tube and sac for a comparative description.
The new species is related to P. frisoni
Banks in sharing the characters of the penis
lacking a caecum, and tergum ten with two
elevated sensilla basiconica patches.
The Holotype will be deposited in the
National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
(USNM). Paratypes will be deposited in the
following museums and individual collec-
tions: Bill P Stark, Clinton, Mississippi
(BPS), C. P. Gillette Museum of Arthropod
Diversity, Colorado State University
(CSU), Monte L. Bean Museum, Brigham
Young University (BYU), and Ralph F
'The views of the author do not purport to reflect
the position of the Department of the Army or the
Department of Defense.
Perlesta, new species, North America, Nearctic Region, Virginia
Kirchner (RFK, Huntington, West Virgin-
1a).
Perlesta teaysia Kirchner and
Kondratieff, new species
(Figs. 1-10)
Male.—Forewing length 7.5-8.5 mm.
General color light yellow in life, pale yel-
low in alcohol. Head with a brown patch
over ocellar triangle, with a light brown
spot near anterior margin (Fig. 1). Prono-
tum brown. Wings pale, costal margin of
forewing pale. Femora with dorsal margin
brown. Tergum 10 with two elevated patch-
es of 19 to 21 brown sensilla basiconica,
separated along mid-line of tergum (Figs.
2, 8, 9). Paraprocts straight, moderately
long without a tooth (Figs. 3, 4). Penis tube
and sac long and slender; dorsal patch pale
and inconspicuous; caecum absent (Fig. 5).
Female.—Forewing length 10-11 mm.
Color pattern similar to male. Subgenital
plate with a prominent U-shaped notch, out-
er lobes rounded (Fig. 6).
Egg.—Length ca. 0.37 mm, width ca.
0.29. Collar short, ca. 0.025 high, ca. 0.073
wide. Chorion smooth (Fig. 10).
Nymph.—Body length 8-10 mm. Gen-
VOLUME 99, NUMBER 2 291
mo N
er
< <>
i ed a) :
ve
. SES
Figs. 1-7. Perlesta teaysia. 1, Adult head and pronotum. 2, Male terminalia, dorsal. 3, Paraproct, lateral. 4,
Paraproct, caudal. 5, Penis, lateral (arrow indicates penis armature). 6, Female subgenital plate, ventral. 7,
Nymphal head and pronotum.
292 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
M
Figs. 8-10. Perlesta teaysia, scanning electron photomicrographs. 8, Male terminalia, dorsal, 372. 9, Sen-
silla basiconica, 10th tergum, male, 1260. 10, Egg, 238.
eral color yellow brown; body clothed with
appressed brown hairs. Head without trans-
verse dark band crossing ocelli (Fig. 7). Oc-
cipital setal row approaches ecdysial suture.
Pronotal disk with “‘parentheses-like”’ dark
brown areas (Fig. 7). Abdominal terga yel-
low brown; intercalary bristles without dark
sockets.
Types.—Holotype d¢, Virginia: Wythe
County, Reed Creek of New River, U.S.
Route 11 bridge, 3.2 km West of Wythe-
ville, 23 July 1996, R. FE Kirchner and B.
C. Kondratieff (USNM). Paratypes: same
data as holotype, 6 3d, 17 2 (CSU); same
data as holotype except, 11 July 1975, R. E
Kirchner 1 6, 7 2 (RFK); same data as
holotype except 21 6,5 2, (reared), 7 July
1979, R. E Kirchner (BPS, BYU, RFK).
Additional material examined.—13
nymphs, same data as holotype except, 7
July 1979, R. E Kirchner.
Diagnosis.—The adult male of P. teaysia
will key to couplet 3 in Stark (1989), ap-
pearing most similar to P. frisoni. Both spe-
cies have conspicuous elevated patches of
sensilla basiconica on tergum 10 (Figs. 2,
8, 9). However, the paraprocts of P. teaysia
are longer and lack a mesal tooth (Figs. 3,
4; see Stark 1989, Fig. 52). The adult fe-
male subgenital plate of P. frisoni is trian-
gular in shape and has a V-shaped notch
(Stark 1989), while the subgenital plate of
P. teaysia is rounded in outline and has a
U-shaped notch (Fig. 6).
Etymology.—tThe specific epithet refers
to the ancient Teays River system of the
VOLUME 99, NUMBER 2
Pliocene. The present Kanawha-New River
drainage in North Carolina, Virginia, and
West Virginia is considered as a remnant of
the upper Teays River. Reed Creek flows
into the New River 22 km East of Wythe-
ville, Virginia.
Remarks.—Stark (1989) reported P. de-
cipiens (Walsh), P. frisoni and P. placida
from Virginia. The distribution of P. frisoni
is apparently limited to the older Appala-
chians Mountains (Blue Ridge Physio-
graphic Province) of North Carolina, Ten-
nessee and Virginia. The following species
of stoneflies have been collected at the type
locality: Allocapnia granulata (Claassen),
A. loshada Ricker, A. nivicola (Fitch), A.
rickeri Frison, Prostoia completa (Walker),
Strophopteryx fasciata (Burmeister), Tae-
niopteryx burksi Ricker and Ross, 7. maura
(Pictet), Acroneuria abnormis (Newman),
293
A. filicis Frison, Neoperia clymene (New-
man), N. occipitalis (Pictet), Paragnetina
media (Walker), Diploperia morgani Kon-
dratieff and Voshell, Helopicus subvarians
(Banks), and /soperla signata (Banks).
ACKNOWLEDGMENT
We thank Bill P. Stark for a review of the
manuscript and providing the SEM photo-
micrographs.
LITERATURE CITED
Poulton, B. C. and K. W. Stewart. 1991. The stoneflies
of the Ozarks and Ouachita Mountains (Plecop-
tera). Memoirs of the American Entomology So-
ciety 38: 1-116.
Stark, B. P. 1989. Perlesta placida (Hagen), an eastern
nearctic species complex (Plecoptera: Perlidae).
Entomologica Scandinavica 20: 263-286.
Stark, B. P. and H. Rhodes. 1997. Perlesta xube, a
new stonefly species from Nebraska (Plecoptera:
Perlidae). Entomological News 108.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 294-304
ADDITIONAL NOTES ON NEARCTIC BIBIO GEOFFROY
(DIPTERA: BIBIONIDAE)
Scott J. FITZGERALD
Department of Entomology, Colorado State University, Fort Collins, (CO) 80523,
USA:
Abstract.—Six new synonyms of Nearctic Bibio are presented and a former, subspecific
synonym is raised to specific status. New synonyms are: Bibio nigripilus Loew = B.
abbreviatus Loew; B. painteri James and B. knowltoni Hardy = B. alexanderi James; B.
nigrifemoratus Hardy = B. atripilosus James; B. imparalis Hardy = B. fluginatus Hardy;
B. utahensis Hardy = B. similis James. Variation within these species is discussed, and
diagnoses and distribution are given for each. Bibio xanthopus palliatus McAtee is raised
to specific status, and the morphological similarity of this species to B. xanthopus Wie-
demann is discussed.
Key Words: Bibio, Nearctic, synonymy
Hardy (1945) provided the most recent
revision of the Nearctic species of Bibio.
Presently, 43 species are recognized (Hardy
1981, Fitzgerald 1996, Fitzgerald and
Skartveit, in press).
A number of species of Bibio are based
soley on leg color and thoracic pile color,
both which can be useful in combination
with other characters. These characters of-
ten exhibit much intraspecific variation.
The use of male terminalia, the number of
sensilla of the hind tibia, relative length of
the anterior spur of the fore tibia, shape of
the hind tibial spur, shape and length of the
hind basitarsus, length of the basal portion
of Rs vein relative to r-m crossvein, and
pigmentation of the posterior veins, in com-
bination with leg and pile color, has allowed
better resolution of intraspecific variation,
thus resulting in the synonyms established
in this paper.
The approximate number of sensilla of
the hind tibia of males and some females is
given for the first time for the Nearctic spe-
cies discussed. Although ranges of sensilla
of some species overlap broadly, others do
not, and so can provide an additional char-
acter to aid in species separation.
Sex ratio in several Palearctic Bibio spe-
cies has been shown to be male biased
(more males/female) (Skartveit 1993). Be-
cause males are more commonly collected
and often provide more characters for spe-
cific separation, female records are included
(under material examined for each species)
only when associated with males or for spe-
cies in which females are distinctive. Fe-
males of a number of species are at present,
difficult, or not possible to distinguish.
Terminology follows McAlpine (1981).
In diagnoses, the number of sensilla of the
hind tibia is given as a range where N =
number of hind tibia examined. Acronyms
for specimen depositories used in this study
are as follows: American Museum of Nat-
ural History (AMNH); Monte L. Bean Life
Science Museum, Brigham Young Univer-
sity (BYU); California Academy of Sci-
ences (CAS); Canadian National Collec-
tion, Ottawa (CNC); C. P. Gillette Museum
VOLUME 99, NUMBER 2 295
5
Figs. 1-6. Male postabdomen, dorsal view. 1, Bibio abbreviatus. 2, B. fraternus. 3, B. alexanderi. 4, B.
fluginatus. 5, B. palliatus. 6, B. xanthopus.
of Arthropod Diversity, Colorado State Uni- Natural History, Smithsonian Institution
versity (CSUFC); University of Colorado (USNM).
(UCoIB); Museum of Comparative Zoology, LAs :
Harvard (MCZ); New York State Museum, Bibio abb 4 eviatus Loew
Albany (NYSM); Snow Entomological Mu- (Fig. 1)
seum, University of Kansas (UKaL); Texas Bibio abbreviatus Loew 1864: 54. Syntype
A & M University (TAMU); University of male (MCZ), USA: District of Columbia,
Minnesota (UMSP); National Museum of Osten Sacken; examined.
296
Bibio nigripilus Loew 1864: 55. Syntype
male (MCZ), CANADA: Winnipeg, Ken-
nicot; examined. NEW SYNONYM.
Bibio lucens Hardy 1937: 204. Holotype
male (UMSP), CANADA: Ontario, Min-
ers Bay, 26 May 1929, G.S. Walley. (syn-
onymized with B. nigripilus by Hardy
1945).
Discussion.—Bibio abbreviatus and B.
nigripilus have historically been separated
by the yellow thoracic pile in B. abbrevia-
tus and black thoracic pile in B. nigripilus
(Loew 1864, Hardy 1945, Hardy 1958).
Hardy (1958) stated, “I am very skeptical
of this character and doubt that it alone
would be reliable.”” Examination of male
terminalia of syntypes of both species has
indicated that B. abbreviatus and B. nigri-
pilus are conspecific. Thoracic pile color
may be yellow, black or intermixed.
Diagnosis.—Bibio abbreviatus is most
similar to B. fraternus Loew with which it
is seasonally and geographically sympatric.
However, males of B. abbreviatus can be
separated from B. fraternus by the shorter
hind basitarsus (see Hardy 1958: 30, Figs.
9b and 32, Fig. 12a), and the gonostylus
slightly flattened (Fig. 1), rather than digi-
tate (Fig. 2). Males of B. abbreviatus can
be distinguished from other Nearctic Bibio
by the combination of the following char-
acters: Legs predominantly yellow orange;
anterior spur of fore tibia three-quarters to
subequal length of posterior spine; posterior
basitarsus not swollen; posterior veins con-
colorous with membrane; hind tibial spurs
slender (rather than broadly flattened); gon-
ostylus slightly flattened (Fig. 1); epandrial
cleft broadly V-shaped extending about
one-half length of epandrium (Fig. 1); sen-
silla of hind tibia number approximately
20—200 (average = 108, N = 22).
Females of B. abbreviatus are most sim-
ilar to B. fraternus but can be distinguished
by the black thorax in B. abbreviatus and
the orange yellow thorax in B. fraternus.
Although females of B. fraternus usually
have the thorax distinctly orange yellow,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
the thorax is occasionally nearly black with
only an orange tinge, and should not be
mistaken for B. abbreviatus. Females of B.
abbreviatus can also be distinguished by
the following combination of characters:
Color of legs, length of anterior spur of fore
tibia, and hind tibial spurs as in male; pos-
terior veins pigmented brown; dorsum of
thorax black; sensilla of hind tibia number
approximately 34—89 (N = 4).
Biology.—The flight period of this spe-
cies is April and May with a few records
from June. One record (UMSP) from Min-
nesota in October is probably incorrect.
Adults have been swept from winter wheat
in Indiana. Strickland (1916) reported the
larvae as a celery pest and provided details
of larval feeding habits and biology.
Distribution.—As in Hardy (1958), wide-
spread in eastern United States and south-
eastern Canada: Ontario and Quebec, south
to Georgia, Arkansas, west to South Da-
kota, Kansas, and Texas.
Material examined.—In addition to the
types, the following material was exam-
ined: CANADA: ONTARIO: Lockburn, 18
May 1926, 2 3 1 @ (one pair in copula)
(CSUFC); USA: ARKANSAS: Crawford
Co., 4 April 1927, 2 ¢ (CSUFC); KAN-
SAS: Douglas Co., 16 April 1946, R.H.
Beamer, 3 6 (UKaL); Gove Co., 20 April
1930, 1 6 1 2 (CSUFC); Reno Co., Hutch-
inson, 2 May 1948, R.H. Beamer, 4 ¢d
(UKaL); INDIANA: Tippecanoe Co., La-
fayette, swept from winter wheat, 2 May, 3
36 (USNM); MAINE: Somerset Co., Hwy
201, 12 June 1993, Kondratieff & Bau-
mann, 1° ¢°1 2 Gn’ copula) (SURG);
MARYLAND: Plummers Island, 26 April
1923, H.S. Barber, 1 6 (USNM); MIN-
NESOTA: Cook Co., Min. ES., Hovland,
Malaise trap, 10 October 1973, 1 6
(UMSP); Pine Co., Mine dump, north bank
Snake River, 4 miles east of Pine City, 19
May 1951, 1 6 (UMSP); Pipestone Co.,
Pipestone National Monument, Malaise
trap, 30 May 1973, 1 d (UMSP); MISSOU-
RI: May, C.V. Riley, 1 ¢6 (BYU); NEW
YORK: Hamilton Co.: 6 miles east of In-
VOLUME 99, NUMBER 2
dian Lake, 45°10'14”, 18 May 1977, 1 ¢
(CSUFC); 10 miles east of Indian Lake,
43.45.30—74.10.14, 555 m, T.L. McCabe:
25 May 1980, 1 ¢ (NYSM); 27 May 1980,
1 36 (NYSM); 18 May 1977, 1 6 (NYSM);
OHIO: Fairfield Co., 2 May 1931, J. Pattan,
1 6 (UMSP); PENNSYLVANIA: Centre
Co., State College, 1 May 1910, 1 ¢
(USNM); TEXAS: Bandera Co., Lost Ma-
ples State Park, 22 March 1985, Kovarik,
Jones & Haack, 12 6 1 & (TAMU); VIR-
GINIA: Fauquier Co., Warrenton, 5 May
1928, L.C. Woodruff, 4 ¢ (UKaL); Fairfax
Com Great Ealls;423: April 1919; “1, 6d
(USNM); Montgomery Co., near Toms
Creek, Route 655, 25 April 1979, B. Kon-
dratiefi, tid. (CSUEC),
Bibio alexanderi James
(Fig. 3)
Bibio alexanderi James 1936: 1. Holotype
female (AMNH), USA: Colorado: Boul-
der Creek bottoms near Valmont, 24
April 1934, E. Gordon Alexander; ex-
amined.
Bibio painteri James 1936: 2. Holotype fe-
male, allotype male (same pin) (AMNH),
USA: Kansas, Manhattan, 19 April 1932,
R.H. Painter; examined. NEW SYNONYM.
Bibio knowltoni Hardy 1937: 202. Holotype
male (BYU), USA: Utah, Granger, on Le-
pidium, 29 April 1931, G.E Knowlton;
not examined. NEW SYNONYM.
Bibio knowltoni var. paltidus Hardy 1937:
203. Holotype male (BYU), USA: Utah,
Provo, 8 May 1937, D.E. Hardy. (syn-
onymized with B. knowltoni by Hardy
1965).
Discussion.—The holotype female, allo-
type male (AMNH) and two female para-
types (CSUFC) of B. alexanderi, the holo-
type female, allotype male (AMNH), and
five paratypes (two pair in copula)
(CSUFC) of B. painteri, four male para-
types (USNM) of B. knowltoni, and six
male paratypes (USNM) of B. k. paltidus
were examined and found to be conspecific.
In contrast to the rounded apex of the gono-
297
stylus in many Nearctic Bibio, the gono-
stylus of B. alexanderi is apically acute in
dorsal view (Fig. 3). Previously, males of
B. alexanderi, B. painteri and B. knowltoni
were separated by minor differences in leg
color and pile color of the eyes and tibiae
(James 1936, Hardy 1945). Females were
separated by the entirely orange yellow dor-
sum of the thorax in B. alexanderi, with
black markings in B. painteri, and entirely
black in B. knowltoni. Individual females
from single populations can exhibit this
range of thoracic color.
Hardy (1945) distinguished B. carri Cur-
ran from B. alexanderi by the position of a
transverse depression separating the upper
and lower portion of the compound eye. In
B. carri this depression is just below middle
line, and in B. alexanderi it is near the low-
er one-fourth. A topotypic male of B. carri
(CSUFC) has been examined and is similar
to B. alexanderi in all respects, including
male terminalia, with the exception of the
position of the transverse depression on the
eye. However, since the two species are ap-
parently allopatric (B. carri known only
from Alberta, Canada), and the unique po-
sition of the transverse depression of the
eye in B. carri is not present in any popu-
lations of B. alexanderi examined, B. al-
exanderi and B. carri are recognized as dis-
tinct species.
Diagnosis.—Males of B. alexanderi are
most similar to B. carri, but can be sepa-
rated by the position of the transverse de-
pression of the compound eye (see discus-
sion). The most diagnostic character of B.
alexanderi is the broadly flattened hind tib-
ial spur which will distinguish both males
and females from the similar B. abbrevia-
tus, B. xanthopus, and B. atripilosus. Ad-
ditional characters to distinguish males of
B. alexanderi are: Legs predominantly yel-
low orange, sometimes with brown mark-
ings; anterior spur of fore tibia one-third to
slightly over one-half length of posterior
spine; hind basitarsus not swollen, shorter
(less than four times as long as wide) than
in B. xanthopus Wiedemann; hind tibial
298
spurs broadly flattened and usually apically
rounded; posterior veins darker than mem-
brane; transverse depression separating the
upper and lower portion of the compound
eye near the lower one-fourth of the eye;
gonostylus apically acute in dorsal view
(Fig. 3); sensilla of hind tibia number ap-
proximately 34—78 (N = 33).
Females of B. alexanderi are diagnosed
by the length of the anterior spur of the fore
tibia and hind tibial spur as in the male;
thoracic color ranges from black to orange
yellow to orange yellow with black mark-
ings; sensilla of hind tibia number approx-
imately 34—56 (N = 6).
Biology.—The flight period is March to
early July, with most records from March
and April. Pairs in copula have been taken
in April in Arizona and Colorado. Adult
collection records are from Lindheimera
texana (Asteraceae) in Texas, and on Med-
icago sativa L. (alfalfa: Fabaceae), Lepidi-
um sp. (Brassicaceae), and Descurainia so-
phia (L.) (Brassicaceae) in Utah. It was re-
ported as a nuisance pest in Denver Co.,
Colorado from a bluegrass lawn in late
April. The use of a sprinkler system appar-
ently triggered a mass emergence of the
flies (10—15 insects per blade of grass) (per-
sonal communication).
Distribution.—Southwestern United
States: Arizona, California, Colorado, Kan-
sas, New Mexico, Oklahoma, Texas, and
Utah. Hardy (1945) listed B. painteri from
Ohio.
Material examined.—In addition to the
type material, the following material was
examined: USA: ARIZONA: Gila Co.,
Globe, 5 April 1937, Parker, 3 ¢ 1 2 (one
pair in copula) (USNM); Maricopa Co.,
Phoenix, 8 March 1945, EH. Parker, 1 3
(USNM); CALIFORNIA: Los Angeles Co.,
Glendora, Dalton Canyon, 15 March 1929,
E.G. Anderson, 1 6 (UMSP); COLORA-
DO: Denver Co., Thorton, lawn, 26 April
1994, C. Wilson, 4 d 3 2 (CSUFC); Yuma
Cov (Chief ‘Creek; roade CC» N25. April
1993, S. Fitzgerald, 2 6 (CSUFC); KAN-
SAS: Sedgwick Co., Wichita, 18 April
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1917, 1 6d (USNM); NEW MEXICO: Dona
Ana Co., Las Cruces, N.M.S.U., 13 March
1994, R. Durfee, 1 ¢ (CSUFC); OKLA-
HOMA: Alfalfa Co., 12 April 1931, 2 6
(AMNH); Cleveland Co., Norman, 13
March 1931, 9 dg 2 2 (AMNH); Stephens
Co., Comanche, 12 April 1952, Michener,
Beamers, Wille & LaBerge,5 622 (1 ¢
taken on Lindheimera texana) (UKaL);
TEXAS: 12 ‘“Mareh -1900;0 1 1@e 22
(USNM); Travis Co., Austin, 9 March
1900, 4 6 4 2 (USNM); UTAH: Grand
Co., Colorado River, highway 163 north of
Moab, 23 April 1982, Baumann & Clark, 3
6 (in alcohol) (BYU); Washington Co.:
North Fork Virgin River, Temple of Sina-
wara, Zion National Park, 30 March 1981,
Baumann & Stranger, 3 d (in alcohol)
(BYU); Emerald Pools, Zion National Park,
8 April 1983, Baumann, 1 46 (in alcohol)
(BYU); Utah Co., Provo, Environs., D.E.
Hardy, 1 3d (BYU).
Bibio atripilosus James
Bibio atripilosus James 1936: 2. Holotype
male (AMNH), USA: Colorado, Boulder,
5 May 1934, M.T. James; examined.
Bibio nigrifemoratus Hardy 1937: 206. Ho-
lotype male and allotype female (CNC),
CANADA: British Columbia, Monte
Lake, 13 May 1936, J.K. Jacob; exam-
ined. NEw SYNONYM.
Bibio nigrifemoratus var. gilvus Hardy
1937: 206. Holotype male (USNM),
USA: Utah, Hyrum, 1 May 1937, G.E
Knowlton, EC. Harmston; examined.
(synonymized with B. nigrifemoratus by
Hardy 1961).
Discussion.—The holotype (AMNH) and
four male paratypes (CSUFC) of B. atripi-
losus, the holotype of B. n. gilvus, and the
holotype, allotype (CNC) and six paratypes
(BYU) of B. nigrifemoratus were examined
and found to be conspecific. The shape of
the gonostylus of the male is distinct from
all other Nearctic Bibio (see Hardy 1961:
185, Fig. 6c). Characters historically used
to separate B. atripilosus and B. nigrife-
VOLUME 99, NUMBER 2
moratus (Hardy 1945), such as thoracic pile
color, wing length, and length of the ante-
rior spur of the fore tibia, vary intraspecif-
ically. Bibio nigrifemoratus was described
with black pile, and B. atripilosus described
with a wing length of 7 mm. In males of
this species thoracic pile color ranges from
yellow to black, the wing length ranges
from 4—6 mm, and the length of the anterior
spur of the fore tibia is one-fifth to one-
third the length of the posterior spine.
Of the two paratypes of B. nigrifemora-
tus that Hardy (1937) listed from Kiger’s
Island, Oregon (Benton Co.), one has been
examined. This specimen is not B. nigrife-
moratus, but B. similis James, differing in
the longer anterior spur of the fore tibia and
the simple, digitate rather than uniquely
shaped gonostylus (see Hardy 1961: 185,
Fig. 6c). The second paratype specimen
from Kiger’s Island, Oregon is also proba-
bly B. similis.
Diagnosis.—Males of B. atripilosus are
most similar to B. xanthopus, but can be
distinguished by the unique shape of the
gonostylus (see Hardy 1961: 185, Fig. 6c).
Males can be distinguished from all other
Nearctic Bibio by the following combina-
tion of characters: Anterior spur of fore tib-
ia short, one-fifth to one-third length of pos-
terior spine; hind basitarsus not swollen;
posterior veins darker than membrane; hind
tibial spurs slender (rather than broadly flat-
tened); sensilla of hind tibia number ap-
proximately 15-50 (N = 13); gonostylus
distinct (see Hardy 1961: 185, Fig. 6c).
Biology.—The flight period is April to
June, with most records in April and May.
Collections have been made at elevations of
610-1830 m.
Swarms of 5—15 males, 0.30—0.61 m off
the ground, have been observed in an open
ponderosa pine forest (elevation ~ 1830 m)
along the Front Range in northern Colora-
do. Swarms were generally oriented over
rocks, and when clouds temporarily
blocked direct sunlight, individuals would
land on rocks or adjacent vegetation. With
the return of direct sun, swarming would
299
resume. Copulating pairs have been ob-
served in April and May. In one case, two
males attempted to copulate, and simulta-
neously displaced each other from one fe-
male.
Distribution.—Northwestern United
States and extreme southwestern Canada:
British Columbia, California, Colorado,
Idaho, Montana, Utah, and Washington.
Material examined.—In addition to type
material, the following material was ex-
amined: CANADA: BRITISH COLUM-
BIA: Robson, 26 May 1936, Foxlee, 2 ¢
(USNM); USA: CALIFORNIA: Cuyamaca,
17 May 1948, 1 d (USNM); COLORADO:
Boulder Co.: 5 May 1934, M.T. James, 1 3
(CSUFC); one mile southwest of Boulder,
6000 ft., 25 April 1963, U.N. Lanham, 3 6
(UCoIB); Boulder, 24 April 1971, U.N.
Lanham, 3 6 (UCoIB); Larimer Co., Hor-
setooth Reservoir, ridge north of Spring
Creek Dam, 24 April 1995, S. Fitzgerald &
A. Foley, 25 6 5 @ (two pairs in copula)
(CSUFC); IDAHO: Boise Co., Warm
Spring Creek junction South Fork Payette
River, highway 21, 23 April 1984, Bau-
mann & Nelson, 1 6 (BYU); UTAH: Car-
bon Co., Minnie Maude Creek junction
Nine Mile Creek, 1 April 1978, Baumann
& Winget, 1 d6 (BYU); WASHINGTON:
Stevens Co., Kettle Falls, 3 May 1912, 3 6
(USNM); Whitman Co.: Pullman, J.A. Hys-
lop, 25 6 (USNM); Pullman, 2000—2500
ft013 sApml 1991). PF) Mclellan; 1 16
(CSUFC).
Bibio fluginatus Hardy
(Fig. 4)
Bibio fluginatus Hardy 1937: 201. Holotype
male (USNM), CANADA: British Co-
lumbia, Salmon Arm, 13 May 1933,
Hugh Leech; examined.
Bibio imparalis Hardy 1959: 209. Holotype
male (CAS), USA: California, Mariposa
Co., Yosemite Valley, 21 May 1921, E.C.
Van Dyke; not examined. NEw SyYN-
ONYM.
Discussion.—The descriptions of B. flu-
ginatus and B. imparalis are nearly identi-
300
Figs. 7—10.
palliatus. 9, B. xanthopus. 10, B. similis.
cal. Neither relationships nor diagnostic
characters were given by Hardy (1959). The
male terminalia of the holotype male of B.
fluginatus and a topotypic male paratype of
B. imparalis (USNM) have been examined
and are found to be conspecific. The holo-
type of B. fluginatus is missing the hind
legs, which have important characters for
species separation, but Hardy (1937) pro-
vided a good original description of the
hind legs.
Diagnosis.—Males of this small species
of Bibio are most similar to B. atripilosus,
but can be distinguished by the simple dig-
itate gonostylus rather than the uniquely
shaped gonostylus of B. atripilosus (see
Hardy 1961: 185, Fig. 6c). Males can also
be distinguished by the following combi-
nation of characters: Legs yellow orange;
anterior spur of fore tibia one-forth to one-
third length of posterior spine; wing 4 mm,
posterior veins darker than membrane; hind
basitarsus slightly swollen; hind tibial spurs
slender; antennal flagellum five segmented;
sensilla of hind tibia number approximately
78—98 (N = 3); in dorsal view gonostylus
broadly rounded apically (Fig. 4); epandrial
cleft about two-thirds length epandrium,
broadly V-shaped, usually with a U-shaped
notch anteriorly (Fig. 4).
Biology.—All known specimens were
collected in May.
Distribution.—Very few specimens are
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ee eee |
Male postabdomen. 7, Bibio similis, dorsal view. 8—10, Right gonostylus, posterior view. 8, B.
available for study. The known range pres-
ently includes California, British Columbia,
and Utah.
Material examined.—Both species were
previously known only from the types. In
addition to the type material listed above,
the following specimen was examined:
USA: UTAH: Summit Co., Yellow Pine
Campground, 27 May 1980, R. Shaha, 1 d
(BYU).
Bibio palliatus McAtee, New Status
(Figs. 5, 8)
Bibio xanthopus palliatus McAtee 1922:
16. Holotype male, allotype female (same
pin) (USNM), USA: Idaho, Moscow, 7
May 1894; examined.
Bibio signatus Hardy 1937: 208. Holotype
male (BYU), USA: Utah, Spanish Fork,
29 April 1936, D.E. Hardy; examined.
(synonymized with B. palliatus by Hardy
1945).
Discussion.—Bibio x. palliatus was orig-
inally described by McAtee (1922) as a
subspecies of B. xanthopus Wiedemann
(1828: 80) for those specimens differing
from ‘‘typical’’ xanthopus by the coloration
of the legs and thoracic pile, and relative
size. Hardy (1937) described B. signatus as
a distinct species, later synonymized it with
B. x. palliatus (Hardy 1945), and eventually
considered both as synonyms of B. xantho-
VOLUME 99, NUMBER 2
pus (Hardy 1965). Examination of the ter-
minalia of male holotypes of B. signatus
and B. x. palliatus, as well as many speci-
mens of B. xanthopus, indicates that B. pal-
liatus should be considered a distinct spe-
cies. Females of the two species are pres-
ently indistinguishable.
Although the type of B. xanthopus is a
female (Hardy 1967), the identity of this
species cannot be mistaken for B. palliatus
due to geographic proximity. Bibio xantho-
pus was described from New York and is
widespread throughout the United States (as
Hardy 1945), whereas B. palliatus is re-
stricted to the northwestern United States
and southwestern Canada.
Hardy (1945) provided some characters
for the separation of the larvae of B. xan-
thopus and B. palliatus.
Diagnosis.—Males of B. palliatus are
most similar to B. xanthopus and B. similis
James, but can be distinguished by the api-
cal portion of the gonostylus very short in
dorsal view (Fig. 5) and the basal portion
of the gonostylus broadly rounded in pos-
terior view (Fig. 8), whereas B. xanthopus
and B. similis have the apical portion of the
gonostylus more elongate in dorsal view
(Fig. 6 and 7) and the basal portion of the
gonostylus more slender in posterior view
(Figs. 9 and 10). Males of B. palliatus can
also be distinguished by the following com-
bination of characters: Anterior spur of fore
tibia one-half to two-thirds (usually just
over one-half) length of posterior spine;
posterior veins darker than membrane; hind
tibial spur slender; gonostylus with distal
portion very short in dorsal view (Fig. 5),
and broadly rounded basally in posterior
view (Fig. 8 and also Hardy 1961: 187, Fig.
7c; this figure is under the name B. utah-
ensis (see discussion of B. similis)); hind
basitarsus not swollen, elongate, robust;
sensilla of hind tibia number approximately
21-88 (N = 24).
Variation.—The leg color is extremely
variable, ranging from orange yellow with
dark joints, to entirely black, to dark brown
to black with base of hind femur yellow, to
301
entirely orange yellow with dark joints and
the apical three-quarters of the hind tibia
brown black and tarsi brown black, to the
hind femur black and the tibia and tarsi
brown orange. Although McAtee (1922)
described B. palliatus as having “‘somewhat
greater average size’’, wing length ranges
from about 5—8 mm, which broadly over-
laps the range of wing length of B. xantho-
pus. The r-m crossvein ranges from one-
third to equal the length of the base of Rs.
Biology.—The flight period is March—
June, with most records in April and May.
Adults have been collected from wetlands
in Utah and along streams, rivers, lakes,
reservoirs, ponds, and springs in other areas
of the West, which would suggest that moist
or saturated soils are the preferred larval
habitat. However, adults have also been col-
lected from sand dunes with vegetation in
Washington. Specimens have been taken at
elevations of 610-762 m in Washington
and approximately 1525 m in Colorado.
Distribution.—Northwestern United
States and southwestern Canada: Alberta,
British Columbia, California, Colorado,
Idaho, Montana, Nevada, Oregon, western
South Dakota, Utah, Washington, and Wy-
oming.
Material examined.—In addition to the
type material, the following material was
examined: CANADA: ALBERTA: Belly
River Campground, Waterton Lakes Na-
tional Park, 22 May 1993, Baumann & Liu,
1 6 (BYU); North Willow Creek, highway
22, 28 May 1993, Baumann & Liu, 3 6
(BYU); USA: CALIFORNIA: Nevada Co.,
Bear River, highway 20, Placer Co. line, 21
April 1987, Baumann, Nelson & Wells, 1
¢ (in alcohol) (BYU); COLORADO: Lar-
imer Co., Dixon Reservoir, 14 May 1994,
B.C. Kondratieff, 2 ¢6 (CSUFC); IDAHO:
Nez Perce Co., Mission Creek, highway 95,
2 miles west of Culde Sac, 27 April 1985,
Baumann & Nelson, | ¢d (BYU); MON-
TANA: Glacier Co., South Fork Cut Bank
Creek, highway 89 north of Kiowa, 22 May
1993, Baumann & Liu, 5 ¢d (BYU); Teton
Co., North Fork Sun River, highway 287
302
north of Augusta, 21 May 1993, Baumann
& Liu, 1 6 (BYU); OREGON: Multnomah
Co., Wahkeena Creek at Wahkeena Falls,
29 March 1984, G.R. Fiala, 1 d (in alcohol)
(BYU); Umatilla Co., Wildhorse Creek at
Athena, 25 April 1985, Baumann & Nel-
son, 3 3 (in alcohol) (BYU); Wallowa Co.,
Rock Creek, highway 82, 3 miles northwest
of Wallowa, 19 May 1977, Baumann &
Dunster, 4 6 8 @ (in alcohol) (BYU);
SOUTH DAKOTA: Custer Co., Iron Creek,
Dakota Lake, 5 June 1995, Baumann &
Huntsman, 1 3d (BYU); Pennington Co.:
spring at Whitewood Creek, 7 June 1995,
Baumann & Huntsman, 4 6 (BYU); Iron
Creek, Spearfish Canyon, 7 June 1995,
Baumann & Huntsman, 1 ¢ (BYU);
UTAH: Washington Co., Santa Clara River,
headwaters, North Juniper Campground, 26
May 1976, Baumann, | ¢ (in alcohol)
(BYU); Utah Co.: Powell Slough, 8 May
1980, S.M. Clark, 4 d (CSUFC); Lehi En-
virons., 4 May 1968, W. Clark, 1 d6 (BYU);
Provo, 26 May 1955, 1 36 (BYU); South
Fork of Provo Canyon, Agnes Hardy, 1 d
(this specimen is part of the paratype series
of B. utahensis Hardy) (BYU); Provo, 26
April 1958, S.K. Taylor, 2 ¢6 (BYU); Go-
shen Pond, 8 May 1973, S.B. Shurtleff, 1
3 (BYU); Goshen Springs, 1 May 1965,
W.M. Tingey, 1 6 (BYU); wetland west of
Utah Valley State College, 4 May 1996,
R.W. Baumann, 14 ¢ 1 2 (BYU), 14 6 1
2 (CSUFC); Provo, Environs, L.E. Perry,
7 3 (BYU); Provo, Environs, 6 May 1969,
1 3d (CSUFC); WASHINGTON: Benton
Co., Hanford Site, sand dunes with vege-
tation near Hanford townsite, T12N R28E
Section 5, 20 April 1994, R.S. Zach, 5 3 1
2 (CSUFC); King Co.: Seattle, 10 April
1933; 1 3% (CSUFC): Fall City, 14 ‘April
1970, D.R. Harris, 1 ¢6 1 2 (CSUFC); Kit-
titas Co., Yakima River, highway 90, Cle
Elum, 6 May 1982, Baumann & Smith, 1
3 1 2 (in alcohol) (BYU); Whitman Co.:
Pullman, April, 2 ¢ (USNM); Pullman,
2000-2500 ft., 20 April 1991, P. McLellan,
1 6 1 2 (CSUFC); Pullman, May 1921, 2
3 (CSUFC); Pullman, 8 May 1931, 1 ¢
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
(CSUFC); WYOMING: Crook Co., Belle
Fourche River at Hulett, 8 June 1995, Bau-
mann & Huntsman, 2 d (BYU); Niobrara
Co., Quinn Creek, highway 20 at Lusk, 2
June 1995, Baumann & Huntsman, 4 d 2
SIE (Bip Gl OD B
Bibio similis James
(Figs. 7, 10)
Bibio similis James 1936: 5. Holotype male
(AMNH), USA: Colorado, Boulder, 5
May 1934, M.T. James; examined.
Bibio utahensis Hardy 1937: 208. Holotype
male (BYU), USA: Utah, Utah Co., Pro-
vo environs., Harry Thomas; examined.
NEw SYNONYM.
Discussion.—The holotype of B. utah-
ensis is conspecific with B. similis. How-
ever, the paratype series of B. utahensis is
composed of two species, B. similis and B.
palliatus, only separable by examination of
male terminalia. Hardy (1961) illustrated
(p. 187, Figs. 7a—7c) the male terminalia
and hind and fore tibia of B. palliatus under
the name B. utahensis.
Diagnosis.—Males of B. similis are most
similar to B. xanthopus and B. palliatus.
Males of B. similis can be distinguished
from B. xanthopus by the longer anterior
spur of the fore tibia, legs usually dark
brown to black, and the more slender base
of the gonostylus in posterior view (com-
pare Figs. 9 and 10). Bibio similis can be
distinguished from B. palliatus only by ex-
amination of male terminalia; the gonosty-
lus of B. palliatus is robust and broadly
rounded basally in posterior view (Fig. 8)
with a very short apical portion in dorsal
view (Fig. 5), whereas the gonostylus of B.
similis is slender basally in posterior view
(Fig. 10) with the apical portion in dorsal
view, not shortened, but developed into a
longer, simple, digitate process with a blunt
apices (Fig. 7). Males of B. similis can also
be distinguished by the following combi-
nation of characters: Legs predominantly
black (see variation below); anterior spur of
fore tibia one-half to three-fourths length of
VOLUME 99, NUMBER 2
posterior spine; posterior veins darker than
membrane; hind basitarsus not swollen;
hind tibial spur slender; sensilla of hind tib-
ia number approximately 21-51 (N = 18);
gonostylus basally slender and distally
elongate (Figs. 7 and 10).
Variation.—In males thoracic pile color
ranges from yellow to black to intermixed,
and the legs range in color from black to
dark brown to the femora blackish and the
tibia and tarsi brown orange.
Although B. similis was originally de-
scribed with the anterior spur of the fore
tibia not being over one-fourth the length
of the posterior spine, a study of the holo-
type, paratypes, and additional material, re-
veals that the anterior spur ranges from one-
half to three-fourths the length of the pos-
terior spine.
Biology.—The flight period is April and
May. It has been taken on flowers of Salix
(Salicaceae) in Colorado. Adults seem to be
most commonly collected along creeks,
ponds, and rivers, and the larvae may prefer
moist or saturated soils.
Distribution.—Northwestern United
States: Colorado, Oregon, Western South
Dakota, Utah, and Washington.
Material examined.—In addition to the
types, the following material was exam-
ined: USA: COLORADO: Boulder Co., Sa-
whill Ponds, 14 May 1994, B. Kondratieff
& R. Durfee, 12 5 6 @ (one pair in copula)
(CSUFC); Delta Co., 5 miles northeast of
Delta off highway 65, 3 May 1987, U.N.
Lanham & M. Weissmann, 1 6 (CSUFC),
6 3d (UCoIB); Douglas Co., 5 miles south-
east of Franktown, Salix flowers, 26 April
1963, U.N. Lanham, 1 ¢ (UCoIB); Garfield
Co., Grizzly Creek, 9 May 1995, W. Cran-
shaw, 4°61 2 (CSUFC); Larimer ’Co., N.
Colorado Nature Center, 25 May 1987,
B.C. Kondratieff, 1 6 (CSUFC); ORE-
GON: Umatilla Co.: Umatilla River at Mis-
sion, 5 miles east of Pendleton, 25 April
1985, Baumann & Nelson, 2 ¢6 (in alcohol)
(BYU); Walla Walla River in Milton-Free-
water, 25 April 1985, Baumann & Nelson,
1 3 (in alcohol) (BYU); Union Co., Phillips
303
Creek, highway 82, Elgin, 19 April 1977,
Baumann, 1 ¢ (in alcohol) (BYU); Wal-
lowa Co., Rock Creek, highway 82, 3 miles
northwest of Wallowa, 19 May 1977, Bau-
mann & Dunster, 1 ¢ (in alcohol) (BYU);
SOUTH DAKOTA: Lawrence Co., Whit-
wood Creek between junction Yellow and
White Tail Creeks (WW-09), 19 May 1981,
Baumann & Furnish, 1 6 (in alcohol)
(BYU); UTAH: Cache Co., Blacksmith
Fork Canyon, 21 May 1983, C.R. Nelson,
1 d (BYU); Utah Co.: East side Utah Lake:
10 36 (BYU); V.M. Tanner, 3 6 (CSUFC);
20, Aprile tos7. se Shurtlert, yd 112
(CSUFC); Wasatch Co., spring tribs. of
Provo River near Hoovers, 14 May 1987,
Nelson & Wells, 1 6 (BYU); WASHING-
TON: Kittitas Co., Teanaway River, high-
way 10, 4 miles south of Cle Elum, 6 May
1982, Baumann & Smith, 5 ¢ (in alcohol)
(BYU).
ACKNOWLEDGMENTS
I sincerely thank Paul H. Arnaud Jr., Cal-
ifornia Academy of Sciences; Richard W.
Baumann, Monte L. Bean Life Science Mu-
seum, Brigham Young University; Leslie C.
Costa, Museum of Comparative Zoology,
Harvard University; Jeff M. Cumming, Ca-
nadian National Collection of Insects, Ot-
tawa; David Grimaldi and Julian Stark,
American Museum of Natural History; and
FE Christian Thompson, Systematic Ento-
mology Laboratory, USDA, for the loan of
types and critical material. My visit to the
United States National Museum of Natural
History was supported by the Samuel Wen-
dell Williston Diptera Research fund. Spe-
cial thanks are also due to Boris C. Kon-
dratieff, C. P. Gillette Museum of Arthro-
pod Diversity, Colorado State University,
for his continual support of my research and
critical review of the manuscript.
LITERATURE CITED
Fitzgerald, S. J. 1996. Notes on Nearctic Bibio (Dip-
tera: Bibionidae). Proceedings of the Entomolog-
ical Society of Washington 98: 50—54.
Fitzgerald, S. J. and J. Skartveit. In press. Holarctic
304
distributions in the genus Bibio (Diptera: Bibion-
idae). Entomologica Scandinavica.
Hardy, D. E. 1937. New Bibionidae from nearctic
America. Proceedings of the Utah Academy of
Sciences, Arts and Letters 14: 199-213.
1945. Revision of Nearctic Bibionidae, in-
cluding Neotropical Plecia and Penthetria (Dip-
tera). The Kansas University Science Bulletin 30:
367-547.
1958. Guide to the Insects of Connecticut.
Part VI. The Diptera or true flies of Connecticut.
Sixth Fascicle. Family Bibionidae. Bulletin of
Connecticut State Geological and Natural History
Survey 87: 5—45.
1959. A New Bibionidae from California
(Diptera). Pan-Pacific Entomologist 35: 209-211.
. 1961. The Bibionidae of California. Bulletin
of the California Insect Survey 6: 179-196.
1965. Family Bibionidae, pp. 191-196. In
Stone, A., C. W. Sabrosky, W. W. Wirth, R. H.
Foote, and J. R. Coulson, eds., A catalog of the
Diptera of America North of Mexico. Agricultural
Handbook 276, 1696 pp.
. 1967. The Types of Bibionidae (Diptera) in
the Naturhistorisches Museum, Wien. Annalen des
Naturhistorischen Museums in Wien 70: 169-181.
. 1981. Bibionidae, pp. 217—222. In McAlpine
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
J. FE, B. V. Peterson, G. E. Shewell, H. J. Teskey,
J. R. Vockeroth, and D. M. Wood, eds., Manual
of Nearctic Diptera. Vol. 1. Agriculture Canada
Monograph 27.
James, M. T. 1936. Some new western Bibionidae
(Diptera). American Museum Novitates 832, 6 pp.
Loew, H. 1864. Americae septentrionalis indigena.
Centuria quinta. Berliner Entomologische Zeit-
schrift 8: 49-104.
McAtee, W. L. 1922. Notes on nearctic bibionid flies.
Proceedings of the United States National Muse-
um (1921) 60(11 [=2406]), 27.
McAlpine, J. F 1981. Morphology and terminology—
adults. [Chapter] 2, pp. 9-63. In McAlpine J. F,
B: V: Peterson; G! E> Shewell He Jz feskeyaniesine
Vockeroth, and D. M. Wood, eds., Manual of Ne-
arctic Diptera. Vol. 1. Agriculture Canada Mono-
graph 27.
Skartveit, J. 1993. The Norwegian Bibionidae (Dip-
tera): distribution, flight periods, swarming and re-
productive behaviour. MS Thesis, University of
Bergen, Norway, 114 pp.
Strickland, E. H. 1916. The March fly (Bibio abbrev-
iatus) in grain fields and as a pest of celery. The
Agricultural Gazette of Canada 3: 600-603.
Wiedemann, C. R. W. 1828. Aussereuropaische zwei-
flugelige Insekten. Vol. 1, xxxii + 608 pp. Hamm.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 305-311
A NEW SPECIES OF MEALYBUG IN THE GENUS PSEUDOCOCCUS
(HOMOPTERA: PSEUDOCOCCIDAE) OF QUARANTINE IMPORTANCE
DouGLASS R. MILLER AND DOUGLAS J. WILLIAMS
(DRM) Systematic Entomology Laboratory, PSI, Agricultural Research Service,
USDA, Bldg. 046, BARC-W, Beltsville, MD 20705, U.S.A.; (DJW) Department of En-
tomology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.
Abstract.—A new species of mealybug, Pseudococcus odermatti, is described that has
been taken in quarantine on Aglaonema, Citrus, and other plants and is of concern as a
pest. Since it is frequently taken in quarantine at US ports of entry, it is important that
the species be described. The adult female and third-instar female are described and
illustrated and are incorporated in a key.
Key Words:
The genus Pseudococcus encompasses
more than 150 species of mealybugs (Ben-
Dov 1994) including several major pests
such as the citrophilus mealybug P. calceo-
lariae (Maskell), the long-tailed mealybug
P. longispinus (Targioni Tozzetti), the grape
mealybug P. maritimus (Ehrhorn), and the
obscure mealybug P. viburni (Signoret)
(hitherto known as P. affinis (Maskell) see
Ben-Dov and Matile-Ferrero 1995). In re-
cent years an undescribed species of Pseu-
dococcus has been intercepted at ports-of-
entry in the United States and Japan and
has impacted the movement of citrus fruit
between Florida and Japan and between the
Bahamas and the United States. It has po-
tential quarantine implications on the World
movement of ornamentals such as Aglaone-
ma (Araceae). The genus Aglaonema con-
tains many species grown as ornamentals in
subtropical areas and in greenhouses.
The purpose of this research is 1) to de-
scribe the morphological characters of this
undescribed species so that they can be
used to differentiate it from similar species,
2) provide data on hosts and distribution,
and 3) incorporate the species in keys to
Mealybugs, Coccoidea, Pseudococcidae, quarantine, citrus, Aglaonema
adult females of Pseudococcus so that it can
be accurately identified.
METHODS
Terminology in the descriptions follows
that of Williams and Watson (1988) and
Gimpel and Miller (1996). The cerarii are
numbered forward from the anal lobe pair
(number 1) to the frontal pair (number 17).
The cerarian formula is given as the cerar-
ian number with the number of conical se-
tae in parentheses. For example, the cerar-
ian formula 1—11(2) means that cerarii
numbers | through 11 each have 2 conical
setae. Measurements and numbers are from
10 specimens when available, and are given
as an average followed by the range in pa-
rentheses. Depositories of specimens are:
The Natural History Museum, London
(BMNH); California Department of Food
and Agriculture, Sacramento (CDA); Flor-
ida State Collection of Arthropods, Gaines-
ville (FSCA); Muséum National d’ Histoire
Naturelle, Paris (MNHN); University of
California, Davis (UCD); National Museum
of Natural History, Beltsville, MD
(USNM).
306
RESULTS
Pseudococcus odermatti Miller and
Williams, new species
Figs. 1; 2
Type data.—The adult female holotype is
the right specimen of two on a slide with
the left label ‘‘Pseudococcus/12/ Costa
Rica/ ex Aglaonema sp./ [V-29-76 leaf/ Mi-
ami 13578/ E. B. Lee/ Balsam” right label
‘*Pseudococcus odermatti Miller/and Wil-
liams/HOLOTYPE” with a map giving the
position of the holotype. This slide is de-
posited in the USNM. In addition there are
paratypes that are deposited in BMNH,
CDA, FSCA, MNHN, UCD, USNM.
Etymology.—The species epithet is giv-
en in recognition of the many contributions
of Douglas M. Odermatt, Entomologist,
Animal and Plant Health Inspection Service
(APHIS), who has not only provided many
specimens of this species, but also has add-
ed significantly to our knowledge of scale
insects through the thousands of identifi-
cations that he has provided for APHIS
identifiers each year.
Adult female. Slide-mounted charac-
ters.—Holotype oval, length 2.6 mm, width
1.6 mm. Paratypes 2.6(1.9—3.1) mm long,
1.5(1.0-1.9) mm wide.
Dorsum with 17 pairs of cerarii, cerarian
formula as follows: Left side 1—9(2), 10(3),
11(2), 12(3), 13—14(2), 15(3), 16(4), 17(3);
paratypes rarely with 1 conical seta in cer-
arii 9 and 10, 3(2—4) conical setae in cer-
arius 12, with 3(2—3) in cerarius 14, with
4(3-5) in cerarius 16, and 3(3—4) in cerar-
ius 17. Cerarius 12 with 4 auxiliary setae
(paratypes with 2(1—4) setae), 18 trilocular
pores (paratypes with 18(12—22) pores),
and 2 discoidal pores (paratypes with 1(0—
3) pores). Multilocular pores absent; triloc-
ular pores evenly scattered over surface;
discoidal pores of 2 sizes, those associated
with oral-rim or oral-collar tubular ducts
small, about %4 diameter of trilocular pore,
those scattered over surface larger, about %
size of trilocular pore. Oral-rim tubular
ducts each with 1(0—2) associated discoidal
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
pores, without closely associated setae, oral
rims present submarginally posterior of
frontal cerarius, absent between cerarii 15
and 16, present between cerarii 13 and 14,
also present near cerarii 4—8, 11, and 12
(paratypes also usually with oral rims near
cerarii 2 and 9, rarely with oral rim near
cerarius 10), also present submedially and
medially, with 17 oral rims on abdomen
(paratypes with 19(16—22) oral rims). Oral-
collar tubular ducts present marginally and
submarginally around perimeter of body,
also present medially and submedially on
anterior abdominal segments, thorax, and
head, often with 1 or 2 associated discoidal
pores, wall of tube heavily sclerotized, giv-
ing bright orange appearance with phase
contrast microscopy. Longest submedial
seta on abdominal segment VII 57p long
(paratypes 66(49-—101) ww); 5 submedial se-
tae on abdominal segment VIII (paratypes
4(2-8) setae), longest seta 52 w long (para-
types 49(37-72) wp).
Anal ring setae 163 w long (paratypes
169(151-195) w); 1.7 times as long as
greatest diameter of ring (paratypes
1.6(1.4—1.9) times).
Venter with multilocular pores in poste-
rior and anterior bands on abdominal seg-
ments V—VIII, in posterior band on abdom-
inal segment IV (paratypes sometimes with
1 or 2 pores near anterior margin of abdom-
inal segment IV, rarely with only 1 or 2
pores near anterior margin of abdominal
segment III), with 6 pores on thorax (para-
types with 7(1—16) pores). Trilocular pores
scattered over surface. Discoidal pores of
same 2 sizes as on dorsum, smaller size as-
sociated with oral rims, larger size scattered
over surface, without a discoidal near eye.
Oral-rim tubular ducts usually with 1 as-
sociated discoidal, without associated setae,
with 5 ducts on submargin from segment II
to cerarius 13 (paratypes with 5(4—12)
ducts), without duct near frontal cerarius.
Oral-collar tubular ducts of 2 sizes, larger
size present near posterior band of multi-
locular pores and on marginal and submar-
ginal areas of abdomen, thorax, and head;
VOLUME 99, NUMBER 2
apm ap
» > bpee\ditan OP on
>of SN Dery
] ae, 4a SF
fa
4
a4 4
307
vg arp
? a7 arp "4 4
Mert Aaa
.
—
PIP De
6
\ PA
go eae
Fig. 1. Adult female Pseudococcus odermatti Miller and Williams. Costa Rica: April 29, 1976, on Agla-
onema sp. leaf, E. B. Lee (taken in quarantine at Miami, FL).
smaller size near setal bases in medial and
submedial areas of abdomen and thorax, 7
oral collars mesad of cerarius 12 (paratypes
10(7—13) ducts), 9 associated with cerarii
10 and 11 (paratypes with 12(8—15) ducts),
22 posterior of eye (paratypes 21(16—33)
ducts), 8 on each side of head (paratypes
4(2-—6) ducts). Setae as follows: 4 cisanal,
longest 79 long (paratypes 71(52—89),);
longest anal-lobe seta 143 long (paratypes
308
160(128—190)); longest seta on trochanter
148py long (paratypes 136(128—163)).
Circulus 168 w wide (paratypes
153(130—170)), divided by intersegmental
fold. Labium 165u long (paratypes
160(148—203)y). Antennae 8-segmented,
490u long (paratypes 431(384—-620)w).
Legs with 95 translucent pores on hind fe-
mur (paratypes 118(43—222) pores); 55
pores on hind tibia (paratypes 95(41—144)
pores). Femur 279w long (paratypes
276(229-—329)); tibia 279% long (para-
types 277(242-353)y); tarsus 114 long
(paratypes 108(99—124)w.). Tibia/tarsus 2.4
(paratypes 2.6(2.4—3.3)). Hind tibia with 30
setae (paratypes 29(26—35) setae).
Notes.—The adult females are variable
in the number of dorsal oral-collar tubular
ducts, the size of the legs, antennae, and
labium, the number of translucent pores on
the hind femur and tibia, and the length of
the dorsal setae. In general, New World
populations often have shorter dorsal setae,
Chinese specimens have more translucent
pores, and Japanese populations have lon-
ger appendages and a larger tibia/tarsus sta-
tistic. However, this variation overlaps suf-
ficiently among specimens collected at var-
ious localities to suggest that there is only
one highly variable species.
Third-instar female. Slide-mounted char-
acters.—Body oval, length 1.3(1.2—1.4)
mm, width 0.8(0.7—0.8) mm.
Dorsum with 17 pairs of cerarii, cerarian
formula as follows: 1—11(2), 12(3 rarely 2),
13—14(2 rarely 3), 15(3 rarely 1 or 2), 16(3
or 4), 17(3 rarely 4). Cerarius 12 with 1(0-—
1) auxiliary setae), 5(3—6) trilocular pores),
without associated discoidal pores. Multi-
locular pores absent; trilocular pores evenly
scattered over surface; discoidal pores of 2
sizes, those associated with oral-rim or oral-
collar tubular ducts small, about %4 diameter
of trilocular pore, those present in medial
areas of abdominal segments VII and VIII
larger, about % size of trilocular pore. Oral-
rim tubular ducts with 1(0—1) associated
discoidal pores, without closely associated
setae, oral rims present submarginally pos-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
terior of frontal cerarius, associated with
cerarli 12, 8, and 5, rarely 1 duct missing
on | side of body, also present submedially
on | or both sides of metathorax with 4(3—
4) oral rims on abdomen, 5(5—6) on thorax
and head. Oral-collar tubular ducts present
marginally and submarginally around pe-
rimeter of body, also present submedially
on anterior abdominal segments, thorax,
and head, often with | or 2 associated dis-
coidal pores, wall of tube heavily sclero-
tized, giving bright orange appearance with
phase contrast microscopy. Longest sub-
medial seta on abdominal segment VII
30(28-—32)y long; 2 submedial setae on ab-
dominal segment VIII, longest seta 23(20-—
27)p long.
Anal ring setae 109(104—116)y long;
1.6(1.5—1.8) times as long as greatest di-
ameter of ring.
Venter without multilocular pores; triloc-
ular pores scattered over surface. Discoidal
pores same as on dorsum, smaller size as-
sociated with oral rims, larger size in sub-
medial line on each side of abdomen. Oral-
rim tubular ducts usually with 1 associated
discoidal, without associated setae, with
2(1—4) ducts on submargin from abdominal
segment II to cerarius 13, without duct near
frontal cerarius. Oral-collar tubular ducts of
2 sizes, larger size on marginal and sub-
marginal areas of abdomen, thorax, and
head; smaller size near setal bases in medial
and submedial areas of thorax, 1(0—1) oral
collars mesad of cerarius, 1(0O—2) associated
with cerarii 10 and 11, 2 posterior of eye,
1(0—1) on each side of head. Setae as fol-
lows: 4 cisanal, longest 38(31—42)p long;
longest seta on trochanter 85(72—91) long.
Circulus 90(74—99)w wide, divided by
intersegmental fold. Labium 112(104—
119). long. Antennae 7-segmented,
270(264—273)y long. Legs without trans-
lucent pores. Femur 144(136—149)y long;
tibia 118(109-124)y long; tarsus 96(96—
99)u long. Tibia/tarsus 1.2(1.1—1.3). Hind
tibia with 13(12—15) setae.
Notes.—The description of the third in-
VOLUME 99, NUMBER 2 309
Fig. 2. Third-instar female Pseudococcus odermatti Miller and Williams. Belize: April 29, 1976, on Agla-
onema sp. leaf, E. B. Lee (taken in quarantine at Miami, FL).
310
star is based on 28 specimens from Belize
on Aglaonema sp.
Discussion.—Pseudococcus odermatti is
most similar to Pseudococcus comstocki
(Kuwana) (see Williams and Granara de
Willink 1992) and P. gilbertensis Beardsley
(see Beardsley 1966). It is easily distin-
guished from them by the absence of trans-
lucent pores on the hind coxae. It also can
be distinguished from P. comstocki by hav-
ing no multilocular pores on the ventral sur-
face of abdominal segment II and from P.
gilbertensis by having more than 5 ventral
oral-collar tubular ducts associated with
cerarius 12. Pseudococcus comstocki has
many multiloculars on abdominal segment
II and P. gilbertensis has fewer than 5 ven-
tral oral-collar tubular ducts associated with
cerarius 12.
The area of origin of this species appears
to be Old World since its closest relatives
seem to be the old world species P. com-
stocki and P. gilbertensis and since the
most common hosts Aglaonema and Citrus
are from the Old World.
Keys.—The key to adult females of
Pseudococcus presented by Williams and
Granara de Willink (1992) should be mod-
ified as follows to accomodate the new spe-
cies:
Change couplet 7 as follows:
7. Oral-rim tubular ducts absent from head near
frontal cerarius; dorsal setae short, less
than 30 long calceolariae (Maskell)
— Oral-rim tubular ducts on 1 or both sides of
head near frontal cerarius; dorsal setae long,
at least 40 long
7A. Oral-rim tubular ducts few, present usually
next to frontal cerarii, and between cerarii 15
and 16, and with | pair near abdominal cer-
arii only; absent from midline of abdomen
and submarginally agavis MacGregor
— Oral-rim tubular ducts relatively abundant,
present usually next to frontal, and near cer-
arii 4—8, 11 and 12, and between 13 and 14;
without oral rim between 15 and 16; present
near midline of abdomen and submarginally
PPS ah we esd odermatti Miller and Williams n.sp.
Specimens Examined.—Paratypes. BA-
HAMAS: Marsh Harbor, Abaco, X-23-92,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
on Citrus paradisi, R. L. Morris (1 ad 2 on
1 sl) BMNH; New Providence, XI-8-78, on
Annona squamosa, C. W. Smith (11 ad &,
2 third instar 2 on 3 sl); no specific locality,
II-13-85, on Aglaonema sp., H. D. Hanna-
gan (taken in quarantine at Miami, FL)(1
ad 2 on 1 sl) USNM; no specific locality,
I-19-86, on Annona squamosa, C. Cohen (1
ad Yon 1 sl); no specific locality, II-7-94,
IX-25-95, on Citrus latifolia, R. Morris
(taken in quarantine at Ft. Pierce, FL)(2 ad
2, 1 third instar 2 on 3 sl) USNM. BE-
LIZE: VIII-13-75, on Aglaonema sp., H. L.
Rubin (taken in quarantine at Miami,
FL)(16 ad 2, 26 third instar 2, 2 second
instar 6 on 6 sl) BMNH, CDA, USNM.
CHINA: Canton, V-18-37, V-17-39, on
Aglaonema sp., (taken in quarantine at
Washington, DC)(3 ad 2 on 2 sl) USNM;
IV-23-40, IV-26-40, on Aglaonema sp.,
(taken in quarantine at Seattle, WA)(2 ad 9
on 2 sl) MNHN, USNM. COSTA RICA:
IV-29-76, on Aglaonema sp. leaf, E. B. Lee
(taken in quarantine at Miami, FL)(4 ad 9
on 4 sl) BMNH, UCD, USNM. HONG
KONG: Sheung Shui, [V-13-93, on Pittos-
porum tobira, C. S. K. Lau (1 ad @ on 1
sl) BMNH; I'V-3-36, V-21-40, on Aglaone-
ma sp., Adams and Limber (5 ad @ on 2
sl) USNM.
UNITED STATES: FLORIDA: Homestead,
IJ-22-73, on Pyracantha sp., W. H. Pierce
(1 ad @ on 1 sl) FSCA; I-7-93, III-1-93,
V-6-93, on grapefruit, T. Sugimoto (taken
in quarantine at Yokohama, Japan)(3 ad @
on 3 sl) BMNH, USNM. HAWAII: VIII-
16-94, on Diospyros blancoi, Hara and
Maemoto (2 ad 2 on 2 sl) USNM.
Other material, not paratypes (all in
USNM unless otherwise stated). CHINA:
IV-26-40, on Aglaonema sp., (taken in
quarantine at Washington, DC)(2 ad @ on
1 sl); V-23-41, [X-23-41, on Aglaonema
sp., (taken in quarantine at Blaine, WA and
Hoboken, NJ)(3 ad 2 on 2 sl). BAHAMAS:
Marsh Harbor, Abaco, X-23-92, on Citrus
paradisi, R. L. Morris (3 ad @ on 3 sl); TI-
8-94, on Citrus latifolia, R. Morris (taken
in quarantine at Ft. Pierce, FL)(1 ad 2on 1
VOLUME 99, NUMBER 2
sl) BMNH. JAPAN: VI-20-33, on Fatsia
japonica, W. H. Wheeler (taken in quaran-
tine at Seattle, WA)(1 ad 2 on 1 sl); [X-13-
35, on Fatsia japonica, K. E. Miller (taken
in quarantine at New Orleans, LA)(2 ad @
on | sl).
SUMMARY
Pseudococcus odermatti occurs on a
number of agricultural and ornamental
hosts and could potentially pose a threat as
an important pest. It is reported on Agla-
onema (Araceae), Annona (Annonaceae),
Citrus (Rutaceae), Diospyros ( Ebenaceae),
Fatsia (Araliaceae), Pittosporum (Pittospo-
raceae), Pyracantha (Rosaceae), and is
known from Florida and Hawaii in the
United States and from the Bahamas, Be-
lize, China, Costa Rica, Hong Kong, and
Japan.
ACKNOWLEDGMENTS
We give special thanks to Paul A. Cour-
neya who is the Animal and Plant Health
Inspection Service (APHIS) Identifier at
Orlando, Florida for helping locate material
of this species and for making many excel-
lent slide preparations. To Linda Lawrence,
Systematic Entomology Laboratory, we
also give thanks for making the exquisite
311
illustration of the adult female. We are
grateful to Douglas M. Odermatt, Entomol-
ogist; Yair Ben-Dov, Institute of Plant Pro-
tection, Volcani Center, Bet Dagan, Israel;
and Michael E. Schauff and Arnold S.
Menke, Systematic Entomology Labora-
tory, USDA, Washington, D.C., for review-
ing the manuscript and making useful crit-
icisms.
LITERATURE CITED
Beardsley, J. W. 1966. Insects of Micronesia Homop-
tera: Coccoidea. Insects of Micronesia 6: 377—
562.
Ben-Dov, Y. 1994. A systematic catalogue of the
mealybugs of the World with data on geographical
distribution, host plants, biology and economic
importance. Intercept Ltd., Andover, UK, 686 pp.
Ben-Dov, Y. and Matile-Ferrero, D. 1995. The iden-
tity of the mealybug taxa described by V.A. Sig-
noret. Bulletin de la Société Entomologique de
France 100: 241-256.
Gimpel, Jr., W. E and Miller, D. R. 1996. Systematic
analysis of the mealybugs in the Pseudococcus
maritimus complex. Contributions on Entomolo-
gy, International 2: 1-163.
Williams, D. J. and Granara de Willink, M. C. 1992.
Mealybugs of Central and South America. CAB
International, Wallingford, UK, 635 pp.
Williams, D. J. and Watson, G. W. 1988. The scale
insects of the tropical South Pacific Region. Part
2. The mealybugs. CAB International, Walling-
ford, UK, 260 pp.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 312-334
SIX NEW SPECIES OF GALL MIDGES (DIPTERA: CECIDOMYITIDAE) FROM
MELALEUCA (MYRTACEAE) IN AUSTRALIA
RAYMOND J. GAGNE, JOSEPH K. BALCIUNAS, AND DAMIEN W. BURROWS
(RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S.
Department of Agriculture, c/o National Museum of Natural History, NHB 168, Wash-
ington, D. C. 20560, USA.; (JKB) USDA-ARS Australian Biological Control Laboratory,
James Cook University, Townsville, 4811, Queensland, Australia; present address: West-
ern Weeds Quarantine Laboratory, Agricultural Research Service, USDA, 800 Buchanan
St., Albany, CA 94710, U.S.A.; (DWB) Australian Centre for Tropical Freshwater Re-
search, Australian Biological Control Laboratory, James Cook University, Townsville,
4811, Queensland, Australia.
Abstract.—A new genus, Lophodiplosis Gagné, is described for five new species of
gall midges from Australia associated with Melaleuca spp. At least two of these are
candidates for biological control of the introduced pest, Melaleuca quinquenervia, or
paperbark, in Florida. In addition, a new species of Lasioptera is described. The new
species are: Lophodiplosis indentata and Lophodiplosis denticulata that form blister galls
on leaves of Melaleuca spp.; Lophodiplosis bidentata, responsible for rosette bud galls
on Melaleuca spp.; Lophodiplosis cornuata, forming trumpet-shaped leaf galls on Me-
laleuca viridiflora; Lophodiplosis trifida, an inquiline in galls of L. indentata, L. denti-
culata, and L. bidentata; and Lasioptera uncinata, an inquiline in galls of L. indentata
and L. cornuata. Keys are given to adults of Australian genera of the supertribe Ceci-
domyiidi and to adults and pupae of the six species found on Melaleuca spp. during this
study.
Key words: gall midges, Cecidomyiidae, Melaleuca, Australia
Melaleuca quinquenervia is one of more
than 200 species of a large genus of Myr-
taceae that is mostly endemic to Australia
(Barlow 1988, Holliday 1989). While most
Melaleuca species are small shrubs, M.
quinquenervia grows into a robust tree up
to 25 m high in Australia (Bodkin 1991)
and up to 29 m high in Florida (Rockwood
and Geary 1991). It is one of many Austra-
lian trees that are popular in plantings in
tropical and subtropical regions of the
world. Unfortunately, M. quinquenervia has
become a pest in some locations in the
United States. It is considered the most
troublesome terrestrial weed in Florida
(Florida Conservation Foundation (1993)
where it was introduced in 1906 (Schmitz
et al. 1991). It is a lesser pest on some Ha-
waiian Islands (Balciunas, unpublished). In
southern Florida this plant now occupies at
least 500,000 acres (Bodle et al. 1994),
causing extensive environmental and eco-
nomic damage (Balciunas and Center
1991). Even small trees produce a great
number of seeds that in Florida, unlike in
Australia, result in a thick carpet of seed-
lings that in a few years form dense, mon-
ospecies forests (Balciunas, personal obser-
vation). In Australia, native insects suppress
the growth of saplings (Balciunas and Bur-
VOLUME 99, NUMBER 2
rows 1993). In Florida, conventional con-
trol measures, such as cutting, burning, and
herbicides, have proven to be ineffective,
costly, or environmentally inappropriate in
managing this pest. Since 1986, a consor-
tium of Florida and U.S. agencies has sup-
ported a U.S. Department of Agriculture
project in Australia to locate, test, and ex-
port potential biological control agents to
help control this weed. The faunal surveys
of M. quinquenervia, the largest plant ever
targeted for a classical biological control ef-
fort (Balciunas et al. 1994), have found
more than 450 herbivorous insects associ-
ated with this tree in Australia (Balciunas
et al. 1995). Included among these insects
are gall midges that form or are associates
of various leaf and bud galls. This paper
treats six species of gall midges found as-
sociated with three kinds of galls (Figs. 1—
6) on M. quinquenervia and some of its
close allies in the Melaleuca leucadendra
group (Blake 1968).
All six species of Cecidomyiidae are new
to science and described here. Four species
are gall makers and the remaining two are
presumed to be inquilines or successors.
The gall midges causing leaf blister and bud
rosette galls show potential as biological
control agents and will be investigated fur-
ther. The leaf blister galls were the most
commonly found in our survey and abun-
dant from August to November with the
emergence of new leaves. Heavily infested
leaves may be rolled or otherwise distorted,
but even small numbers of galls are a tax
on the host’s energy. The rosette bud galls
appear to have a more deleterious effect on
Melaleuca quinquenervia by killing branch
terminals.
METHODS AND MATERIALS
Surveying, collecting, and rearing for
this study in Australia were done by the
staff of the Australian Biological Control
Laboratory under the direction of JKB and
DWB. Galls were collected on members of
the Melaleuca leucadendra complex mainly
in coastal areas of Queensland between
S)IL3}
Townsville (19°28’S) and Cairns (16°54’S).
Some collections were also made in the vi-
cinity of Brisbane (27°30’S) in southern
Queensland and of Cooktown (15°28’S) in
northern Queensland. Plant species sampled
were: Melaleuca arcana S.7T. Blake, M.
dealbata S.T. Blake, M. ‘fluviatilis’? (Bar-
low 1988), M. leucadendra (L.) L., M.
quinquenervia (Cav.) S.T. Blake, M. saligna
Schauer, and M. viridiflora Sol. ex Gaertner.
Galls from the field were either preserved
in alcohol or kept in small plastic containers
that were checked daily for emergence. Lar-
vae and pupae were excised from sample
galls, adults were reared and their pupal ex-
uviae saved, and all specimens were pre-
served in 70% ethanol. Some larvae and
adults were mounted for microscopic study
in Canada balsam, using the method out-
lined in Gagné (1989a, 1994), some were
prepared for SEM viewing. In the following
descriptions, anatomical terminology of the
adult stage follows McAlpine (1981) and
that of the larval stage follows Gagné
(1989a). Specimens used in this study are
deposited in the Australian National Insect
Collection, Canberra (ANIC) or the U.S.
National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
(USNM). Other abbreviations used in the
text are for collectors in the lists of speci-
mens studied: ADM = A.D. Moore; DWB
= D.W. Burrows; JKB = J.K. Balciunas;
RJG = R.J. Gagné.
Lophodiplosis Gagné, new genus
Adult.—Head: Eyes large, connate, eye
bridge about 12 facets long; facets hexag-
onal, closely adjacent throughout. Vertex
with short to long dorsal protuberance bear-
ing 2—6 setae. Frons with 6—12 setae. La-
bella elongate-hemispherical in frontal
view, the apices incurved and acute, each
with 5—12 setae. Palpus 4 segmented. An-
tenna with 12 flagellomeres, first and sec-
ond flagellomeres connate, apex of twelfth
flagellomere with narrow, elongate exten-
sion. Male flagellomeres binodal, some-
times weakly so, with 1 circumfilum on ba-
314
sal node, 2 on distal node, circumfila sep-
arate or running into one another, the loops
short to long. Female flagellomeres becom-
ing successively slightly shorter from base
to apex of antenna, nodes parallel sided or
constricted near middle, necks prominent
and without setulae; circumfila made up of
two wavy horizontal bands connected by 2
vertical bands, closely embracing node for
all or most of extent.
Thorax: Scutum with 2 lateral and 2 dor-
socentral rows of setae interspersed with
sparse scales. Mesanepisternum covered
with scales on dorsal half to two thirds. Me-
sepimeron with vertical row of setae. Wing:
R,; curved apically to join C posterior to
wing apex; C broken beyond juncture with
R;; Rs apparent as spur of R;, variable in
position; Cu forked; M,,, present as fold.
Tarsal claws curved beyond midlength,
one- or two-toothed or simple; empodia al-
most reaching bend in claws; pulvilli about
one fourth length of empodia.
Male abdomen: First through sixth ter-
gites entire, rectangular, with posterior row
of setae, usually with lateral setae, anterior
pair of trichoid sensilla, and otherwise
mostly covered with setiform scales. Sev-
enth and eighth tergites more weakly scler-
otized; seventh tergite usually with 1—sev-
eral posterolateral setae, a few scales, with
anterior pair of trichoid sensilla; eighth ter-
gite bare except for anterior pair of trichoid
sensilla. Cerci variable, triangular or ovoid.
Hypoproct bilobed, variable in shape and
vestiture. Aedeagus short to moderately
long. Gonocoxite with small, obtuse me-
sobasal lobe. Gonostylus tapered gradually
from wide base to narrower, toothed apex,
with scattered setae, setulose basally, re-
mainder of surface marked with minute lon-
gitudinal ridges.
Female abdomen: First through seventh
tergites entire, rectangular, with posterior
row or rows of setae, usually with lateral
group of setae, anterior pair of trichoid sen-
silla, and otherwise covered with setiform
scales; eighth tergite shorter, narrower, and
more weakly sclerotized than preceding,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
with either weak, scattered setae or strong
setae mixed with scales posteriorly and an-
terior pair of trichoid sensilla; ovipositor
short-protrusible, less than 1% length of
seventh tergite, with ventral setae on inter-
segmental membrane, evenly distributed se-
tae on ninth segment, with or without setae
on tenth tergum; cerci short to long, more
or less bilaterally flattened, with 2—4 ven-
troapical, thick, setiform sensoria on each,
and otherwise covered with setulae; hypo-
proct variably shaped.
Pupa.—Vertex with conspicuous projec-
tion. Two pairs of vertexal papillae present,
one of each pair with elongate seta and en-
larged base. Antennal bases evenly rounded
or angular. Face with or without horns, with
or without pair of setae mesally. Prothorac-
ic spiracle several times as long as wide.
Abdominal segments uniformly spiculose,
without spines.
Larva.—Third instar: Body flattened-cy-
lindrical, rounded at both ends. Integument
mostly rugose. Antenna less than twice as
long as wide. Spatula with two acutely tri-
angular anterior projections, intervening
concavity smooth to minutely dentate. Pap-
illar pattern generally as for Cecidomyiidi
(Gagné 1989a): lateral papillae on each side
of spatula reduced in some species to the
two setose pairs; dorsal and pleural papillae
with setae little longer than wide; terminal
segment with 2 or 4 pairs of papillae.
Second instar: Spatula present. Other-
wise as for third instar.
Type species.—Lophodiplosis indentata
Gagné.
Etymology.—Lophodiplosis is of femi-
nine gender and combines the Greek “‘lo-
phos,’ meaning “crest”? with “‘diplosis.”
The prefix refers to the protruding vertex of
the pupa. The suffix has been commonly
used for genera of the supertribe Cecido-
myiidi and means “‘double,”’ with reference
to the binodal male flagellomeres.
Relationships.—Lophodiplosis belongs
to the supertribe Cecidomyiidi. Uniquely
derived characters (sometimes secondarily
lost) of the supertribe are the dorsal protu-
VOLUME 99, NUMBER 2
berance of the adult head and the binodal,
tricircumfilar male antennal flagellomeres.
This genus differs from all other known
genera of Cecidomyliidi except Acacidiplos-
is in the development of the pupal vertex
into a protuberance or crest instead of or in
addition to the antennal horns. This devel-
opment appears to be another strategy to
allow pupae to cut their way out of the
galls. Although the five species placed here
are otherwise diverse, we regard the pupal
head character as a shared, derived char-
acter until we have better knowledge of the
Australian cecidomyiid fauna. The presence
of a similar modification in Acacidiplosis,
an African genus restricted to Acacia spp.,
presumably arose separately in that genus
because not all Acacidiplosis species have
it (Gagné and Marohasy 1994) and the two
genera share no other obvious derived char-
acters. Acacidiplosis differs from Lophodi-
plosis in having no vertexal protuberance in
the adult, less than four palpal segments,
and small, asetulose, dorsoventrally com-
pressed and mesally joined female cerci.
KEY TO ADULTS OF AUSTRALIAN GENERA
OF THE SUPERTRIBE CECIDOMYIIDI
Lophodiplosis is the first genus of Ceci-
domyiidi outside of the tribe Asphondyliini
to be based exclusively on Australian spe-
cies. Except for the Skuse species, which
remain essentially unknown (Gagné
1989b), all the non-asphondyliine Cecido-
myliidi identified from Australia to date are
satisfactorily placed in cosmopolitan genera
(Gagné 1989b). The nine genera keyed be-
low are doubtless only a small sample of
what will be found in Australia, but the key
serves to place Lophodiplosis for future
identification. Following each generic name
is the most up-to-date and comprehensive
reference. For a list of Australian species in
each of the genera, see Gagné (1989b).
1. Male gonostylus situated on dorsum of gono-
coxite; female seventh sternite at least 1%
times length of sixth sternite
— Male gonostylus situated on apex of gonocox-
ite; female seventh sternite not appreciably lon-
SersthanesiXthipewea se een Ney, ees 4
2. Empodia much shorter than tarsal claws; male
flagellomeres with necks; female without dor-
sal pair of lobes at base of intersegmental
MeEMbranevOmovipositon eam... ee
Skusemyia (see Kolesik 1995a)
— Empodia as long as tarsal claws; maie flagel-
lomeres without necks; female with dorsal pair
of lobes at base of intersegmental membrane
of ovipositor
3. Surface of flagellomeres covered with closely
anastomozing circumfila; ovipositor short, pli-
able Eocincticornia (see Kolesik 1995b)
— Surface of flagellomeres with only weakly an-
astomozing circumfila in male but not in fe-
male; ovipositor long, needle-like, sclerotized
Asphondylia (see Gagné 1994)
4. Tarsal claws curved near basal third ....... 5
— Tarsal claws curved beyond midlength
5. Head without vertexal protuberance
shag tits, EAR eee Diadiplosis (see Gagné 1994)
— Head with vertexal protuberance
Se eee eee aa Ee Feltiella (see Gagné 1995)
6. Head with vertexal protuberance; male flagel-
lomeres with 3 circumfila occasionally inter-
connected Lophodiplosis
— Head without vertexal protuberance; male fla-
gellomeres with 2 circumfila
7. Palpus 3 segmented, third segment inserted
subapically on second; female cerci bilaterally
compressed . Zeuxidiplosis (see Harris 1966)
— Palpus 3 or 4 segmented, if with only 3, the
third segment inserted apically on second; fe-
male cerci dorsoventrally compressed ...... 8
8. Abdominal second through seventh tergites
without lateral setae at midlength ......
ha Slade vet woee ees Dae Stenodiplosis (see Gagné 1994)
— Abdominal tergites with lateral setae at mid-
lensth ieee. ce Contarinia (see Harris 1979)
THE SPECIES OF LOPHODIPLOSIS
Besides the five new species keyed and
described below, two Australian species re-
ported earlier from Eucalyptus, a genus
close to Melaleuca, may also belong to Lo-
Phodiplosis. These are ‘‘Cecidomyia”’ par-
ilis Skuse (1888: 87) and ‘“‘Cecidomyia”’
eucalypti Skuse (1890: 381), both described
on the basis of superficial characters. ““Ce-
cidomyia”’ parilis was originally described
from a female collected by Skuse from
Middle Harbour, New South Wales. This
pinned specimen, with abdomen and ap-
pendages intact, is in the ANIC and labeled,
316
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-6. Galls of Lophodiplosis spp. on Melaleuca spp. Figs. 1—2, Blister leaf galls of L. indentata on M.
dealbata: 1, 1x; 2, 6x, leaf in cross section showing larval feeding chamber and exit tube after adult has emerged.
Figs. 3—4, Rosette bud gall of L. bidentata on M. quinquevervia: 3, 1x; 4, 6x, group of larval cells with adjacent
modified leaves from center of bud, one cell entire, three with apex dehisced following successful departure of
adult (one in x-section), and one with hole made by parasitoid. Figs. S—6, Trumpet leaf galls of L. cornuata on
M. viridiflora; 5, 1x; 6, 6x, two galls, one in cross section.
“Dipl. parilis, 2 type, EA.A. Skuse, Mid-
dle Harb., S.”’ Skuse (1890) later identified
as this species females bred from leaf blis-
ter galls on Eucalyptus corymbosa collected
by Froggatt at Waverley, near Sydney. The
leaf blisters suggest those of L. indentata
on Melaleuca. Four female specimens im-
mediately follow the type of C. parilis in
the ANIC, each bearing a single label read-
ing, “bred Nov. 20.” These may be Frog-
gatt’s specimens, but until these and the
type are slide mounted, we cannot know
whether Froggatt’s specimens really belong
to “C.” parilis or even if “Cy “parilisais
in Lophodiplosis. *‘Cecidomyia”’ eucalypti
known from evidently a single female
(“‘Description drawn from fresh speci-
men’’) from Botany, New South Wales, was
reared from woody swellings on stems of
Eucalyptus haemastoma. There is no spec-
imen labelled eucalypti in the Skuse collec-
tion ANIC.
VOLUME 99, NUMBER 2
Another Australian species, *‘Cecido-
myia” frauenfeldi Schiner (1868: 7) was
reared from rosette-like malformations cov-
ering the swollen branch buds of a species
of Melaleuca found at ‘‘Naraby Lagoon”
[prob. for Narabeen Lagoon per note by
Skuse 1888: 62], Port Jackson [Sydney,
New South Wales]. The adults were de-
scribed as having more than 12 flagellom-
eres, with those of the male stemmed,
which indicates they belong to neither Lo-
phodiplosis nor Lasioptera, the two genera
we have found on Melaleuca.
KEY TO ADULTS OF LOPHIDIPLOSIS SPP. AND
LASIOPTERA ON MELALEUCA
1. R; about half wing length (Fig. 58); inquiline
or successor in blister galls of Lophodiplosis
indentata and trumpet galls of Lophodiplosis
CONMUAIOV Roy eeo. Sic ey: Lasioptera uncinata
— R, reaching costa beyond wing apex (Fig. 11)
Set neal Mee aM apis, oy Gat ee: sinc ess) Bayesian sive 2
2. Tarsal claws simple (Fig. 10); male flagello-
meres with short, indistinct internodes and cir-
cumfilar loops barely longer than wide (Fig. 8)
Petras coor sA uss yl wea tan 8 «ewan, «ee, oe 3
— Tarsal claws toothed (Fig. 24); male flagello-
meres with distinct internodes and circumfilar
loops much longer than wide (Fig. 21) ..... 4
3. Tarsal claws amber colored; vertexal protuber-
ance elongate with apical and lateral setae (Fig.
7); male circumfila interconnected (Fig. 8);
from leaf blister galls .. Lophodiplosis indentata
— Tarsal claws dark brown; vertexal protuberance
short with only apical setae (Fig. 23); male cir-
cumfila not interconnected (as in Fig. 21); from
trumpet leaf galls Lophodiplosis cornuata
4. Tarsal claws dark brown, with 2 teeth (Fig. 24);
female eighth tergite with distinct row of setae
posteriorly; male cerci triangular (Fig. 25);
from bud rosette galls .. Lophodiplosis bidentata
— Tarsal claws amber colored, with single tooth
(Fig. 18); female eighth tergite with only scat-
tered setae posteriorly; male cerci quadrate or
secondarily lobed! (Figs: 205 41))) 5.5525. ..- 5
5. Vertexal protuberance elongate with lateral se-
tae (as in Fig. 7); aedeagus shorter than hy-
poproct (Fig. 20); setae at base of female cerci
longer than cerci (as in Fig. 13); from leaf blis-
tengcall Spee aes Lophodiplosis denticulata
— Vertexal protuberance short with only apical
setae (as in Fig. 23); aedeagus longer than hy-
poproct (Fig. 41); setae at base of female cerci
much shorter than cerci (Fig. 43); possible in-
quiline in galls of Lophodiplosis indentata, Lo-
Siliy/
phodiplosis denticulata, and Lophodiplosis bi-
dentata
Lophediplosis trifida
KEY TO PUPAE OF LOPHODIPLOSIS SPP
LASIOPTERA ON MELALEUCA
AND
1. Vertex convex, without protuberance (Fig. 64)
se cch, RAGA oe Pe ARR Reeth bates Lasioptera uncinata
Vertex with protuberance (Figs. 44-53) .... 2
. Frons without pair of setae anterior to labrum
(Figs. 44, 46); vertex with | central and 2
much shorter ventral protuberances (Figs. 45,
AT) erat A ae ROR LO Be ete Gees 3
— Frons with pair of setae anterior to labrum; ver-
tex with a single median protuberance that may
be divided apically (Figs. 48, 50, 52)
3. Vertex with central protuberance tapering to
point (Fig. 44) Lophodiplosis indentata
— Vertex with central protuberance cylindrical,
concavesapicallya(hien46) te ae ee
Sti, <Mobaynptere tana Lophodiplosis denticulata
4. Vertex with protuberance shorter than height of
antennallbasesm (alone oan.) leaner een
S SREE oN One ORGS Sys aes Lophodiplosis bidentata
— Vertex with protuberance much longer than
height of antennal bases (Figs. 50, 52)
5. Vertex with protuberance conical from base to
slightly notched apex (Fig. 50)
Se gitinc hac: ME tee ee ME. Lophodiplosis cornuata
— Vertex with protuberance strongly angled for
most of length, with three apical points (Fig.
SD ie nes < eS RNeee sec. coe Lophodiplosis trifida
N |
Lophodiplosis indentata Gagné,
new species
Figs. 1, 2, 7-17, 44, 45
Adult.—Head (Fig. 7): Vertex with elon-
gate dorsal protuberance bearing 4—5 setae,
2-3 at or near apex, the remainder near
midlength. Male flagellomeres (Fig. 8) bin-
odal, internodes very short, necks moder-
ately long, nodes with sparse setulae; cir-
cumfila interconnected, bases not on same
horizontal plane, loops short. Female fla-
gellomeres (Fig. 9) cylindrical with mod-
erately long necks; circumfila appressed ex-
cept for short loops at apex.
Thorax: Wing (Fig. 11), length, 1.9—2.3
mm in males (n = 10), 2.5—3.1 mm in fe-
males (n = 10); Rs apparent only as spur
of R,, situated closer to arculus than to apex
of R,. Tarsal claws (Fig. 10) amber colored,
untoothed.
Male abdomen: First through sixth ter-
318 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
14 15 2s
Figs. 7-15. Lophodiplosis indentata. 7, Male head. 8, Male third flagellomere. 9, Female third flagellomere.
10, Tarsal claw and empodium. 11, Wing. 12, Female postabdomen, seventh segment to end, lateral. 13, Female
cerci, detail. 14, Aedeagus and hypoproct, ventral. 15, Male genitalia, dorsal.
VOLUME 99, NUMBER 2
gites with mostly single, posterior row of
setae, partly double on sixth tergite, with O—
5 lateral setae, and otherwise mostly cov-
ered with scattered setiform scales. Seventh
tergite with O—5 dorsolateral setae, O—sev-
eral setiform scales mesally, and an anterior
pair of trichoid sensilla. Eighth tergite bare
except for anterior pair of trichoid sensilla.
Genitalia (Figs. 14—15): cerci more or less
acute-triangular, with several setae posteri-
orly; hypoproct bilobed, laterally curving
toward venter and nestling aedeagus, with
1-2 pairs of weak setae on posterolateral
margins; aedeagus short, cylindrical, taper-
ing slightly to rounded apex, reaching ap-
proximately to base of concavity of hypo-
proct; gonostylus with setulae present only
at base.
Female abdomen (Figs. 12-13): First
through fifth tergites with mesally single to
laterally double, uninterrupted, posterior
row of setae, sixth and seventh tergites with
mesally double to laterally triple row of se-
tae; third through seventh tergites with in-
creasing number of lateral setae, 3—4 on
third to 15—20 on seventh; first through sev-
enth tergites covered with setiform scales;
eighth tergite with weak, scattered setae
posteriorly and anterior pair of trichoid sen-
silla; ovipositor short-protrusible, distal half
about 14 times as long as seventh tergite,
with short setae ventrally on intersegmental
membrane, ninth segment with evenly dis-
tributed setae; tenth tergum without setae;
cerci short, bilaterally flattened, closely ap-
proximated mesally with several setae, one
pair of setiform sensoria, and otherwise
covered with setulae; hypoproct entire, lon-
ger than wide.
Pupa (Figs. 44—45).—Vertex with one
dorsoventrally flattened, pyramidal projec-
tion and two shorter, ventral projections.
Antennal bases with sharp, transverse, ven-
tral crest. Face without setae or conspicuous
protuberance.
Larva.—Third instar (Figs. 16—17): In-
tegument mostly rugose. Spatula with two
acutely triangular anterior projections, in-
tervening concavity smooth to minutely
319
dentate, the shaft narrowest at midlength,
broadening but less sclerotized posteriorly
and ending abruptly at broad base. Lateral
papillae on each side of spatula reduced in
most specimens to the two setose pairs, oc-
casionally one or both asetose papillae pres-
ent; dorsal and pleural papillae with setae
little longer than wide; terminal papillae re-
duced to two pairs, with setae slightly lon-
ger than wide; anal papillae absent.
Second instar (based on specimen with
fully developed third instar inside): Body
shape as for third instar but shorter. Other-
wise as for third instar including spatula of
similar shape.
Holotype.—d, from blister leaf galls of
M. quinquenervia, Australia, Queensland,
Tully Heads Road #3, 15.3 km ESE Tully,
17-VH-1995, JKB, FSNQMqn95137, de-
posited in ANIC.
Paratypes.—All specimens from blister
leaf galls, Queensland, Australia, most de-
posited in ANIC, with representatives de-
posited in the USNM. From M. quinque-
nervia: 22, Forrest Beach, 16 km SE In-
gham, 6-VII-1987, JKB, NQMqn87073; 1
pupal exuviae, Forrest Beach Swamp, 15.6
km ESE Ingham, 4-X-1994, DWB,
FSNQMgqn94136; 4 larvae, Forrest Beach
Swamp, 15.6 km ESE Ingham, 10-V-1994,
J.R. Makinson, FSNQMqn94058; 6, Card-
well Swamp Site, 2 km SE Cardwell,
31-VIII-1992, em. 8-IX-1992, ADM,
FSNQMgqn92056; 3 larvae, pupa, Edge
Hill, Woodward Park, 4.2 km WNW Cairns
GPO, 14-VIII-1994, P. Geyson, NQMqn940
80; 32, Centenary Park, Cairns, 3-VIII-
1987, JKB, NQMgqn87085; larva, Tully
Heads Rd) #37133" km ESE Tully PO,
27-V1I-1994, JKB, FSNQMqn94094; 5 lar-
vae, Tully Heads Rd #3, 15.3 km ESE Tully
PO, 2-VIII-1994, DWB, FSNQMgqn941 14;
43,32, 2 pupal exuviae, Tully Heads Road
#3, 15.3 km ESE Tully, 17-VHU-1995, JKB,
FSNQMgn95137 (19 more @ and 7 p. exuv.
in alc.; from same lot as L. denticulata); 1
pupal exuviae, Tully Heads Road #3, 15.3
km ESE Tully, 14-VIII-1995, JKB,
FBNQMgqn95163; 4 larvae, Murrigal Cul-
320
(0) 8)
Figs. 16-21.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
20
16-17, Lophodiplosis indentata. 16, Larval spatula and associated papillae. 17, Larval eighth
and terminal segments. 18-21, Lophodiplosis denticulata. 18, Tarsal claw and empodium. 19, Male genitalia,
dorsal. 20, Aedeagus and hypoproct, ventral. 21, Male third flagellomere.
vert, 14.8 km S Tully, 4-X-1994, DWB,
FSNQMgqn94137; 6 larvae, Murrigal Gra-
vel Pit, 19.3 km S Tully, 2-VIII-1994,
DWB, FSNQMgqn94112; larva, Bruce Hwy,
S of Murrigal, 16 km S Tully, 10-II-1992,
ADM, NQMqn92017; 5 larvae, Feluga Site
1, 9.4 km E Tully, 27-VI-1994, JKB,
FSNQMqn94092; larva, Bilyana, Double
Barrel Creek Swamp, 22 km NE Tully,
10-11-1992, ADM, NQMgqn92018; 3. lar-
vae, 3 pupae, Townsville, 21-I]-1995, RJG;
3 larvae, Feluga Site 1, 9.4 km E Tully,
26-X-1994, JKB, FSNQMqn94142; 4 lar-
vae, 4 pupae, South Queensland, XI-1995,
DWB,; 3 larvae, Doolandella, 16 km SSW
Brisbane GPO, 30-V-1994, M. Purcell.
From M. dealbata: 143, Cardwell North, 3
kms N Cardwell, 31-VIII-1992, em. 7-25-
VII-1992, ADM, DWB, FSNQMd192057; 2
larvae, Murrigal Culvert, 14.8km S. Tully,
18-IX-1995, DWB, FSNQMd195188; 264,
2, 2 pupal exuviae, Tully Heads Road #4,
16.9 km ESE Tully, 17-VII-1995, JKB,
FSNQMd195135; 2 larvae, Tully Heads
Road #4, 16.9 km ESE Tully, 27-III-1995,
JKB, FSNQMd195076; 2d, 2, 4 pupal ex-
uviae, Murray River Swamp, 12 km S Tul-
ly, 14-VIII-1995, JKB, FSQNMd195159;
VOLUME 99, NUMBER 2
3, 32, 4 pupal exuviae, Forrest Beach
School, 16 km ESE Ingham, 14-VIII-1995
JKB, FSQNMd195154. From M. viridiflora:
3 larvae, Murrigal section of Bruce Hwy,
19.3 km S Tully, 2-VIII-1994, DWB,
FSNQMvr94115; 32, 1 pupal exuviae, Fe-
luga Site 1, 9.4 km E Tully, 18-VI-1995,
JKB, FSNQMvr95140; 3, 22, 2 pupal ex-
uviae, Feluga Site 1, 9.4 km E. Tully,
15-VIII-1995, DWB, FSNQMvr95165 (in
same lot as specimens of L. denticulata).
From M. arcana: 2 larvae, 1 pupal exuviae,
Cape Flattery Swamp, 41.1 km N Cook-
town, 25-VII-1994, JKB, NQMac94074.
From M. fluviatilis: 3 larvae, pupa, Charters
Towers, 24-II-1995, RJG. From M. saligna:
larvae, Flying Fox Swamp, 57.2 km WNW
Cooktown, 27-VII-1994, JKB, FSNQMsI9
4109.
Etymology.—The name indentata is Lat-
in for “‘untoothed,’’ with reference to the
simple, untoothed tarsal claws of this spe-
cies.
Gall (Figs. 1—2).—This species forms a
leaf blister gall, a convex, occasionally low-
conical swelling that shows equally on both
surfaces of the leaf. Leaves infested with
these galls may be curled and otherwise
distorted, especially when the leaf surfaced
is completely covered with the galls. One
or more larvae may be found in a gall but
each larva is in a discrete cell. As the larva
matures, the tissue above one end of the
gall changes in nature to form a cylindrical
passage extending from just outside the lar-
val cell to the abaxial surface epidermis.
Pupation occurs in the larval cell. The full-
grown pupa pushes its way through a thin,
circular cap that develops on the larval cell
wall, then proceeds through the tunnel and
breaks through the thin leaf epidermis
forming the cover of the tunnel and where
the pupa lodges itself. The adult then breaks
out of the pupa, usually leaving the pupal
exuviae caught part way out of the gall exit.
Remarks.—Lophodiplosis indentata was
commonly reared from leaf blister galls on
several species of Melaleuca in Queens-
land, twice in association with L. denticu-
321
lata (q.v.). The gall surface is rounded ex-
cept on M. dealbata where it is usually
pointed in the center (Figs. 1—2).
This species shares with L. denticulata an
exceptionally long adult vertexa! projection,
the male hypoproct partially enfolding the
aedeagus, and the lack of pupal facial setae.
It differs from L. denticulata in having un-
toothed tarsal claws, very short flagello-
mere internodes and shorter, interconnected
circumfilar loops, longer male gonopods,
and a tapering, pyramidal protuberance on
the pupal vertex.
Lophodiplosis denticulata Gagné,
new species
Figs. 18-21, 46, 47
Adult.—Head (as for Fig. 7): Vertex with
elongate dorsal protuberance bearing 4—5
setae, 2—3 at or near apex, the remainder
near midlength. Male flagellomeres (Fig.
21) binodal, internodes and necks moder-
ately long, nodes setulose; circumfila dis-
crete, bases of each on same _ horizontal
plane, loops moderately and uniformly
long. Female flagellomeres (as in Fig. 9)
cylindrical with moderately long necks; cir-
cumfila appressed except for short loops at
apex.
Thorax: Wing (as in Fig. 11), length,
1.8—2.3 mm in males (n = 3), 2.2—2.3 mm
in females (n = 2); Rs apparent only as a
spur off R;, situated closer to arculus than
to apex of R,. Tarsal claws (Fig. 18) amber
colored, with fine basal tooth.
Male abdomen: First through sixth ter-
gites with mostly single, posterior row of
setae, sometimes mostly double on sixth
tergite, with O—5 lateral setae, and other-
wise mostly covered with scattered setiform
scales. Seventh tergite with O dorsolateral
setae and O-several setiform scales mesally.
Eighth tergite bare except for anterior pair
of trichoid sensilla. Genitalia (Figs. 19—20):
cerci more or less quadrate, longest at pos-
terolateral angle, with several setae poste-
riorly; hypoproct bilobed, laterally curving
toward venter and nestling the aedeagus,
with 1—2 pairs of weak setae on posterolat-
322,
eral margins; aedeagus short, cylindrical,
tapering slightly to rounded apex, reaching
almost to posterior margins of hypoproct;
gonostylus with setulae present only at
base.
Female abdomen: First through fifth ter-
gites with mesally single to laterally double,
uninterrupted, posterior row of setae, sixth
and seventh with mesally double to laterally
triple row of setae; third through seventh
tergites with 3—7 lateral setae; first through
seventh tergites otherwise covered with se-
tiform scales; eighth tergite with weak,
scattered setae posteriorly and anterior pair
of trichoid sensilla; ovipositor similar to
that of L. indentata, short-protrusible, distal
half about 1% times as long as seventh ter-
gite, with short setae present ventrally on
intersegmental membrane, evenly distrib-
uted setae on ninth segment; bare of setae
on tenth tergum; cerci short, bilaterally flat-
tened, closely approximated mesally with
several setae, each with one pair of setiform
sensoria, and otherwise covered with setu-
lae; hypoproct entire, longer than wide.
Pupa (Figs. 46—47).—Vertex with one
elongate, cylindrical, ventromesal projec-
tion with concave, crenulate apex, and two
shorter, ventral projections. Antennal bases
with sharp, transverse, ventral crest. Face
without setae or conspicuous protuberance.
Larva.—Unknown. If similar to L. inden-
tata, specimens of L. denticulata may be
included among those listed for L. inden-
tata.
Holotype.—<d, from blister leaf galls of
M. quinquenervia, Australia, Queensland,
Tully Heads Road site #3, 15.3 km ESE
Tully, 17-VII-1995, JKB, FSNQMgqn95137,
deposited in ANIC.
Paratypes.—All specimens from Queens-
land, Australia, and from blister galls on
Melaleuca spp., deposited in ANIC except
for one male and three pupal exuviae in
USNM: 6, same data as holotype (both
from same lot as L. indentata); 2°, M. quin-
quenervia, Edge Hill, Pease St. Park,
Cairns, 2-IX-1992, em. by 18-IX-1992,
JKB, NQMgqn92060; 6, M. quinquenervia,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Feluga Site 3, 13 km NE Tully, 31-VIII-
1992, em. 6-IX-1992, ADM, FSNQMgqn
92058; 2, 5 pupal exuviae, M. viridiflora,
Feluga Site 1, 9.4 km E. Tully, 15-VIII-
1995, DWB, FSNQMvr95165 (in same lot
as specimens of L. indentata).
Etymology.—The name denticulata is a
Latin diminutive for “‘toothed,” with ref-
erence to the thin tooth present at the base
of the tarsal claws of this species.
Gall.—This species was reared from the
swollen leaf blister galls on M. quinquener-
via and M. viridiflora (Figs. 1—2) similar to
those attributed here mainly to L. indentata.
Both species may cause the galls but L.
denticulata was much less commonly
reared.
Remarks.—This species shares with L.
indentata the exceptionally long adult ver-
texal projection, the male hypoproct par-
tially enfolding the aedeagus, and the lack
of pupal facial setae. It differs from L. in-
dentata in having toothed tarsal claws, lon-
ger flagellomere internodes and longer, dis-
crete, circumfilar loops, shortened male
gonopods, and a longer, more cylindrical
protuberance on the pupal vertex.
This species was reared four times from
leaf blister galls similar to those of L. in-
dentata, from four separate localities in
northern Queensland and represented by
one or two adults each and, in one of those
cases, by five pupal exuviae. In two of
those collections, L. denticulata was found
in association with L. indentata. Larvae, if
similar to those of L. indentata, may be
mistakenly listed under that species. We can
offer no speculation for why these closely
related species occupy similar galls on the
same hosts and at the same sites.
Lophodiplosis bidentata Gagné,
new species
Figs. 3, 4, 22-30, 48, 49
Adult.—Head: Vertex with short dorsal
protuberance bearing 2 setae at or near apex
(Fig. 23). Male flagellomeres (similar to
that of L. denticulata, Fig. 21) binodal, in-
ternodes and necks moderately long; cir-
VOLUME 99, NUMBER 2 323
30
Figs. 22-30. Lophodiplosis bidentata. 22, Female third flagellomere. 23, Vertexal projection. 24, Tarsal claw
and empodium. 25, Male genitalia, dorsal. 26, Aedeagus and hypoproct, ventral. 27, Female postabdomen,
seventh segment to end, lateral. 28, Female ninth segment to end, detail. 29, Larval eighth and terminal segments.
30, Larval spatula and associated papillae.
cumfila discrete, bases of each on same hor- with nodes slightly constricted near basal
izontal plane, loops moderately and uni- third, necks elongate.
formly long. Female flagellomeres (Fig. 22) Thorax: Wing length, 2.4—2.6 mm in
324
males (n = 4), 3.0—3.2 mm in females (n
= 4); Rs situated closer to arculus than to
apex of R,. Tarsal claws (Fig. 24) dark
brown, with two basal teeth, the proximal
tooth finer.
Male abdomen: First through sixth ter-
gites with mostly single, but double later-
ally, posterior row of setae, mostly double
row on sixth tergite, all with 3—5 lateral se-
tae. Seventh tergite with 3—5 dorsolateral
setae, and O-3 scales posterolaterally.
Eighth tergite bare except for anterior pair
of trichoid sensilla. Genitalia (Figs. 25—26):
cerci triangular, laterally curved ventrally,
with several setae along margin; hypoproct
bilobed, with several scattered pairs of setae
ventrally; aedeagus longer than hypoproct,
tapering gradually from base to midlength
and widening slightly beyond to rounded
apex, with rows of several papillae; gon-
ostylus with setulae present on basal fourth.
Female abdomen (Figs. 27—28): First
through fifth tergites with mostly double
posterior row of setae, sixth and seventh
with triple row of setae; third through sev-
enth tergites with increasing number of lat-
eral setae, 3—4 on third to 10 on seventh;
eighth tergite with a mostly double poste-
rior row of strong setae mixed with setiform
scales and anterior pair of trichoid sensilla;
Ovipositor short-protrusible, distal half
about % times as long as seventh tergite,
with many setae present laterally and ven-
trally on intersegmental membrane and ven-
trally on ninth segment; tenth tergum cov-
ered laterally with setae; cerci ovoid, bilat-
erally flattened, closely approximated
mesally, with several setae, each with 3—4
setiform sensoria, and otherwise covered
with setulae; hypoproct broad, deeply bi-
lobed.
Pupa (Figs. 48—49).—Vertex with short,
ventral, triangular, dorsoventrally flattened,
slightly notched projection. Antennal bases
simply rounded. Face with pair of setae an-
terior to labrum, without protuberance.
Larva.—Third instar (Figs. 29-30): In-
tegument mostly smooth. Spatula with two
acutely triangular anterior projections, the
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
shaft narrowest at midlength, broadening
posteriorly and ending abruptly at broad
base. Lateral papillae in two groups of three
on each side of spatula, 2 of each group
with seta, the other without; dorsal and
pleural papillae with setae not longer than
papilla base; terminal segment with 4 pa-
pillae on each side, the setae all tiny but
one on each side slightly longer than re-
mainder.
Second instar: Body shape as for third
instar but shorter. Otherwise as for third in-
star except spatula smaller and not sclero-
tized posteriorly.
Holotype.—d, from rosette galls of M.
quinquenervia, Australia, Queensland, Fe-
luga site #1, 9.4 km E Tully, 15-VIII-1995,
DWB, FSNQMgqn95166, deposited in
ANIC.
Paratypes.—All specimens from M.
quinquenervia, Queensland, Australia, most
deposited in ANIC, with representatives de-
posited in USNM: 26, 32, 2 pupal exuvi-
ae, Feluga site #1, 9.4 km E Tully, 15-VIII-
1995, DWB, FSNQMgqn95166; 2 larvae,
Feluga Site #1, 9.4 km E Tully, 27-VI-
1994, JKB, FSNQMqn94042; pupa, larva,
Feluga Site #1, 9.4 km E Tully, 26-X-1994,
JKB, FSNQMqn94142; 24,12, Feluga site
#1, 9.4 km E Tully, 18-VII-1995, JKB,
FSNQMgqn95139; 1 pupa, Feluga site #1,
9.4 km E Tully, 10-XII-1993, DWB,
FSNQMgqn93084; 2 larvae, Tully Heads
Road #3, 15.3 km ESE Tully PO, 27-VI-
1994, JKB, FSNQMqn94094; ¢, Tully
Heads Road #3, 15.3 km ESE Tully PO,
22-II-1995, RJG; pupa, Murrigal Gravel
Pit, 19.3 km S Tully, 28-III-1995, JKB,
FBNQMgqn95086; 3 pupae, Murrigal Sec-
tion of Bruce Hwy, 19.3 km S Tully, 11-V-
1993, JKB, NQMgqn93025; pupa, larva, 16
km ESE Ingham, Forrest Beach Swamp,
15.6 km ESE Ingham, 2-VIII-1994, DWB,
FSNQMgqn941 10.
Etymology.—The name bidentata is Lat-
in for “two toothed,” with reference to the
two teeth present at the base of the tarsal
claws of this species.
Gall (Figs. 3—4).—The gall made by this
VOLUME 99, NUMBER 2
species is made up of a hard, ovoid larval
cell closely surrounded by modified leaves.
Many separate galls may occur in aggregate
on a single bud. Galls develop on terminal
or lateral buds and cause a foreshortening
of the branch so that all the leaves around
the gall form a rosette. These galls stunt the
surrounding leaves and prevent normal
branch elongation where they occur. Pupa-
tion occurs in the gall. At that time a cir-
cular, weakened area develops at the apex
of the cell. The fully developed pupa push-
es about half way through the trap door
where it lodges. The adult then breaks out
of the pupa.
Remarks.—Lophodiplosis bidentata is
distinct from the other species of Lophodi-
plosis in having two teeth at the base of
each tarsal claw, triangular male cerci and
narrow hypoproct, a short ovipositor with
more than a pair of setiform sensoria on the
cerci and a two-lobed hypoproct, and a
short, dorsoventrally compressed protuber-
ance on the pupal vertex.
Lophodiplosis cornuata Gagné,
new species
Figs: 5516, 31-38, 50,51
Adult.—Head: Vertex with short dorsal
protuberance bearing 2 apical or subapical
setae, placed at or near apex. Male flagel-
lomeres (Fig. 31) only weakly binodal with
very short internode but long neck; | cir-
cumfilum on basal node, 2 on distal node,
all with very short loops. Female flagello-
meres (Fig. 32) cylindrical with moderately
long necks; circumfila appressed.
Thorax: Wing (Fig. 33), length, 2.6—2.8
mm in males (n = 4), 2.8-3.2 mm in fe-
males (n = 4); Rs apparent only as spur of
R;, situated closer to apex of R, than to ar-
culus. Tarsal claws (Fig. 34) dark brown,
untoothed; empodia almost reaching bend
in claws.
Male abdomen: First through sixth ter-
gites with mostly single, sometimes double
laterally, posterior row of setae, sometimes
mostly double row on sixth tergite, with 3—
6 lateral setae. Seventh tergite with 3—6
325
dorsolateral setae, several setiform scales
mesally. Eighth tergite bare except for an-
terior pair of trichoid sensilla. Genitalia
(Figs. 36—37): cerci rounded to blunt pos-
teriorly, with several prominent lateral and
mesal setae; hypoproct deeply lobed, bulg-
ing ventrally, more or less nestling aede-
agus, with several pairs of weak setae ven-
trally; aedeagus shorter than hypoproct, cy-
lindrical, tapering from base to narrow
apex; gonostylus with setulae present on
basal half.
Female abdomen: First through fifth ter-
gites with mesally single to laterally double,
uninterrupted, posterior row of setae, sixth
and seventh with mesally double to laterally
triple row of setae; third through seventh
tergites with increasing number of lateral
setae, 3—4 on third to 15—20 on seventh;
eighth tergite shorter, narrower, and more
weakly sclerotized than preceding, with
scattered weak posterior setae and anterior
pair of trichoid sensilla; ovipositor short-
protrusible, distal half about 1% times as
long as seventh tergite, with short setae
ventrally on intersegmental membrane,
evenly distributed setae on ninth segment;
tenth tergum without vestiture; cerci (Fig.
35) short, bilaterally flattened, closely ap-
proximated mesally with several setae,
those at base no longer than cerci, each with
one pair of setiform sensoria, and otherwise
covered with setulae; hypoproct as wide as
long.
Pupa (Figs. 50—51).—Vertex conical,
more bulbous basally, tapering near mid-
length to dorsoventrally flattened, apically
notched and slightly recurved apex. Anten-
nal bases simply rounded. Face with pair of
setae mesally and without conspicuous pro-
tuberance.
Larva.—Third instar: Integument com-
pletely rugose. Spatula (Fig. 38) with two
acutely triangular anterior projections, the
shaft very short. Lateral papillae in two
groups of three on each side of spatula, 2
of each group with setae, the third without;
dorsal and pleural papillae with setae not
longer than papilla bases; terminal segment
326 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 31-43. 31-38, Lophodiplosis cornuata. 31, Male third flagellomere. 32, Female third flagellomere. 33,
Wing. 34, Tarsal claw and empodium. 35, Female apex of ninth abdominal segment and cerci, lateral. 36, Male
genitalia, dorsal. 37, Aedeagus and hypoproct, ventral. 38, Larval spatula and associated papillae. 39-43, Lo-
phodiplosis trifida. 39, Female third flagellomere. 40, Tarsal claw and empodium. 41, Male genitalia, dorsal. 42,
Female postabdomen, seventh segment to end, lateral. 43, Female apex of ninth abdominal segment and cerci,
lateral.
VOLUME 99, NUMBER 2
with undetermined number of convex pa-
pillae with short setae.
Holotype.—<¢, from trumpet-shaped leaf
galls of M. viridiflora Australia, Queens-
land, Magnetic I., 5-III-1995, RJG, depos-
ited in ANIC. _Paratypes.—AlIl specimens
from trumpet-shaped leaf galls, Queens-
land, Australia, most deposited in ANIC,
with representatives deposited in USNM:
33d, 32, 3 pupal exuviae, on M. viridiflora,
Magnetic I., 5-III-1995, RJG; 22, 7 pupal
exuviae, on M. viridiflora, 16 km ESE In-
gham, Forrest Beach School, 14-VIII-1995,
JKB, FSNQMvr94154; 2 larvae, M. nerv-
osa, Angus Smith Drive, Townsville,
21-HI-1995, DWB.
Etymology.—The name cornuata is Lat-
in for “‘hornlike’’ with reference to the
shape of the gall that suggests the end of a
trumpet.
Gall (Figs. 5—6).—This species forms a
trumpet shaped gall on the abaxial leaf sur-
face of M. viridiflora and M. nervosa. Pu-
pation occurs in the gall. The pupa breaks
out of the apex of the gall and becomes
lodged in the aperture, whereupon the adult
emerges from the pupa.
Remarks.—This species is distinct from
the other species of Lophodiplosis in the in-
conspicuous internode and short circumfila
of the male antenna, the simple, dark brown
tarsal claws, the shape of the male genitalia,
and the large, slightly recurved, conical ver-
texal pupal protuberance.
Lophodiplosis trifida Gagné, new species
Figs. 39-43, 52, 53
Adult.-Head: Vertex with short dorsal
protuberance bearing 2 setae near apex.
Male flagellomeres (as in Fig. 21) binodal,
internodes and necks moderately long,
nodes setulose; circumfila discrete, bases of
each on same horizontal plane, loops mod-
erately and uniformly long but not reaching
base of next node or circumfilum. Female
flagellomeres (Fig. 39) cylindrical with
moderately long necks; circumfila ap-
pressed.
Thorax: Wing (as in Fig. 11), length, 2.0
327
mm in males (n = 1), 2.2—2.3 mm in fe-
males (n = 5); Rs apparent only as spur of
R,, situated closer to arculus than to apex
of R,. Tarsal claws (Fig. 40) amber colored,
toothed.
Male abdomen: First through sixth ter-
gites with mostly single, uninterrupted, pos-
terior row of setae, and 2—6 lateral setae.
Seventh tergite with 2—5 dorsolateral setae
and several setiform scales. Eighth tergite
bare except for anterior pair of trichoid sen-
silla. Genitalia (Fig. 41): cerci barely divid-
ed mesally, with short mesoposterior lobe
and longer more or less triangular latero-
posterior lobe, both lobes setose; hypoproct
deeply lobed, the two lobes narrow and
elongate, setose posteriorly; aedeagus elon-
gate, longer than hypoproct lobes, tapering
gradually from base to blunt apex; gonos-
tylus with setulae present on basal third.
Female abdomen (Figs. 42—43): First
through fifth tergites with mesally single to
laterally double, uninterrupted, posterior
row of setae, sixth and seventh with mesal-
ly double to laterally triple row of setae;
third through seventh tergites with increas-
ing number of lateral setae, 3—4 on third to
15—20 on seventh; eighth tergite with weak
posterior setae and anterior pair of trichoid
sensilla; ovipositor short-protrusible, distal
half about 14 times as long as seventh ter-
gite, with scattered setae ventrally on inter-
segmental membrane; setae evenly distrib-
uted on ninth segment; tenth tergum with-
out setae; cerci elongate-ovoid, each with
one pair of setiform sonsoria; hypoproct en-
tire, longer than wide.
Pupa (Figs. 52—53).—Vertex with elon-
gate mesal projection, deeply angled along
entire length, three-pointed apically. Anten-
nal bases each with ventral, conical horn.
Face with two horns anteriorly near base of
antennae and with pair of setae near middle.
Larva.—Unknown.
Holotype.—<é, from blister leaf galls of
M. quinquenervia, Australia, Queensland,
Tully Heads Road #3, 15.3 km ESE Tully,
17-VII-1995, JKB, FSNQMgqn95137, de-
posited in ANIC.
328 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
; iss 7
aa
Figs. 44-47. Pupae or pupal exuviae of Lophodiplosis spp. 44, L. indentata, ventral. 45, Same, lateral. 46,
L. denticulata, ventral. 47, Same, lateral.
VOLUME 99, NUMBER 2 329
Figs. 48-51. Pupae or pupal exuviae of Lophodiplosis spp. 48, L. bidentata, ventral. 49, Same, lateral. 50,
L. cornuata, ventral. 51, Same, lateral.
330
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 52-53.
Paratypes.—All specimens from Queens-
land, Australia, and from blister galls or ro-
sette bud galls on Melaleuca spp., deposited
in ANIC except for one male and 2 pupal
exuviae in USNM: <4, same data as holo-
type (both from same lot as L. indentata
and L. denticulata); 42, 2 pupal exuviae,
blister galls on M. dealbata, Tully Heads
Road #4, 16.9 km ESE Tully, 17-VI-1995,
JKB, FSNQMd195135 (from same lot as L.
indentata); 3, 2 2°, 2 pupal exuviae, from
rosette galls on M. quinquenervia, Feluga
site 1, 9.4 km E Tully, 18-VII-1995, JKB,
FSNQMgqn95139 (in same lot as L. biden-
tata).
Etymology.—The name trifida is Latin
for three-cleft, with reference to the three-
pointed projection of the pupal vertex.
Gall.—This species is presumably an in-
quiline and was reared from leaf blister
galls with L. indentata and L. denticulata
on M. quinquenervia and M. dealbata, and
Pupal exuviae of Lophodiplosis trifida. 52, Ventral. 53, Same, lateral.
from bud rosette galls with L. bidentata on
M. quinquenervia.
Remarks.—Lophodiplosis trifida is dis-
tinct from the other species of Lophodi-
plosis for the secondarily divided lobes of
the male cerci, the deeply divided male hy-
poproct, and the large antennal and facial
horns and long, three-cleft vertexal protu-
berance of the pupa.
Lasioptera uncinata Gagné, new species
Figs. 54—64
Adult.—Head (Fig. 56—57): Antenna
with pedicel slightly wider than flagellom-
eres, first and second flagellomeres connate;
male with 16-19 flagellomeres, slightly
longer than wide; female (Fig. 57) with 31—
33 flagellomeres, wider than long. Palpus 4
segmented. Labella elongate ovoid, with
strong lateral setae.
Thorax: Wing (Fig. 58), 1.3—1.5 mm
long; R; 0.50—0.54 length of wing (n = 5).
VOLUME 99, NUMBER 2
Figs. 54—57.
to end. 56, Head. 57, Part of flagellum.
Scutum with two lateral and two dorsocen-
tral rows of setae, otherwise completely
covered with scales; scutellum with group
of setae on each side, elsewhere covered
with scales; anepisternum with scales on
dorsal two-thirds; mesepimeron with row of
setae and covered with scales; katepister-
num with scales in center. Tarsal claws with
large basal tooth; empodia as long as claws;
pulvilli about % as long as empodia.
Male abdomen: First through seventh
sclerites rectangular with single, uninter-
rupted row of setae, a pair of anterior trich-
oid sensilla, and covered with scales; eighth
tergite developed only on anterior half,
without posterior sclerotization and setae,
and otherwise as for preceding tergite ex-
cept smaller. Genitalia (Figs. 59—60): gon-
ocoxite cylindrical; gonostylus abrubtly ta-
pered beyond base, setulose on bulbous
base, ridged beyond; cerci convex posteri-
orly, with ventral and apical setae; hypo-
proct wide, convex posteriorly, with apical
Lasioptera uncinata. 54, Head, thorax, and abdomen. 55, Female postabdomen, eighth segment
setae; parameres narrow and tapering be-
yond anterodorsal lobe; aedeagus narrow,
as long as gonocoxite.
Female abdomen (Figs. 54-55): First
through seventh tergites generally as for
male but seventh somewhat smaller in ex-
tent; eighth tergum with 2 discrete, elongate
sclerites, each with trichoid sensilla near
anterior third and several setae posteriorly.
Eighth segment with setation only on ter-
gum; intersegmental membrane with lateral
group of elbowed setae on each side, with
no other setation; ninth segment and cerci
about 4.25 times as long as sixth tergite,
with setae only posteroventrally and poster-
olaterally; fused cerci with dorsolateral, gla-
brous, saddle-shaped area bearing wide,
straight setae anteriorly and two postero-
dorsal rows of uncinate setae continuing to
slightly hooked apex of cerci; remaining,
setulose area of cerci with fine, thin ventral
setae ventrally and thicker, curved, poste-
rior setae.
332
°o
61
Figs. 58—64.
2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
/
‘
59 60
——————
Lasioptera uncinata. 58, Wing. 59, Male genitalia, dorsal. 60, Parameres and aedeagus, dorsal.
61, Larval spatula and associated papillae. 62, Eighth and terminal segments of larva, dorsal. 63, Pupal antennal
horns and vertex, ventral. 64, Pupal head and prothoracic spiracle, lateral.
Pupa (Figs. 63—64).—Vertex concave,
the 2 setae elongate. Antennal bases pointed
and bilaterally flattened apicoventrally.
Face without prominences or setae.
Larva tentatively associated with this
species (Figs. 61—62).—Spatula large, es-
pecially area anteriad of lateral papillae, tri-
dentate anteriorly, the middle tooth the lon-
gest and notched. Sternal and ventral papil-
lae of prothorax without setae; with a group
of 4 lateral papillae on each side of spatu-
lae, 2 of each group with short seta. Ter-
minal segment with 3 pairs of papillae, se-
tae of one pair twice as long as remaining
two.
Holotype.—, from blister galls on M.
dealbata, Australia, Queensland, 3 km N
Cardwell, 31-VIII-1992, em. IX-1992,
DWB, NQMdl192057, deposited in ANIC.
Paratypes.—All specimens from leaf
blister or trumpet-shaped leaf galls on Me-
laleuca spp., Queensland, Australia, most
deposited in ANIC, with representatives de-
posited in USNM: 2<¢, from blister galls on
M. dealbata, 3 km N Cardwell, 31 VIII-
1992, em. I[X-1992, DWB, NQMd192057;
23 from blister galls M. quinquenervia,
blister galls, Cardwell Swamp Site, 2 km
SE Cardwell, 31-VIII-1992, emerged
14-IX-1992, ADM, FSNQMgqn92056; 6,
blister galls, M. quinquenervia, Feluga Site
1, 31-VII-1992, em. 3-IX-1992, Mqn920
59, ADM; 2°, 4 pupal exuviae, from trum-
pet-shaped leaf galls, M. viridiflora, 16 km
ESE Ingham, Forrest Beach School,
14-VIII-1995, JKB, FSNQMvr94153; 2°,
from trumpet-shaped leaf galls, M. viridiflo-
ra, Magnetic I., Townsville, 5-III-1995,
VOLUME 99, NUMBER 2
RJG; 2 larvae, from swollen stem, M. leu-
cadendra, Oonoonba McPherson St.,
Townsville, 5-V-1994, L.M. Brown, 94049.
Etymology.—The name uncinata is Latin
for “hooked,” with reference to the hooked
setae on the female cerci. These hooks are
characteristic of all Lasioptera species ex-
cept for a few species in which they are
secondarily lost.
Gall.—This species may be an inquiline
or a successor. The latter term is used for
species that live in a gall after the gall mak-
er has flown. At least one species of La-
sioptera is known to have that habit (Yu-
kawa and Haitsuka 1994). Lasioptera un-
cinata was reared from leaf blister galls of
Lophodiplosis indentata on M. quinquener-
via and M. dealbata and from trumpet leaf
galls of Lophodiplosis cornuata on M. vir-
idiflora.
Remarks.—The larva referred to this
species is only tentatively placed here, but
fits Lasioptera and most other Lasiopterini
with its four papillae on each side of the
spatula.
Two Australian species of Lasioptera
were described earlier from Melaleuca and
Eucalyptus. Lasioptera nodosae Skuse
(1888:130) was based on a female bred
from deformed buds of Melaleuca nodosa
found in November by Masters in Home-
bush, NSW, and Lasioptera miscella Skuse
(1890: 388) bred from malformed, coales-
cent leaf-stalks of Eucalyptus haemastoma
in November and collected in Botany,
NSW. Peter Kolesik of the University of
Adelaide, who is undertaking a revision of
the Skuse Cecidomyiidae, kindly compared
a photo of the cerci of L. uncinata with
those of females of both Skuse species and
has determined for us that the hooked setae
of the new species are distinct in the num-
ber and position from those of the other
species.
ACKNOWLEDGMENTS
The Australian survey was funded by:
USDA-ARS Office of International Re-
search Programs; U.S. Army Corps of En-
33318)
gineers (Jacksonville District); U.S. Nation-
al Park Service; Florida Department of En-
vironmental Protection; South Florida
Water Management District; Dade County,
Florida; and Lee County, Florida. We are
grateful to Leanne Brown, Jeff Makinson,
and Andrew Moore for assistance in col-
lecting galls in Australia, Peter Kolesik for
his taxonomic help, Nit Malikul for prepar-
ing the microscopic slides, Linda Heath
Lawrence for figures 1—6, and Keith M.
Harris, Kenneth L. Hibbard, Peter Kolesik,
John C. Moser, Allen L. Norrbom, and Nor-
man E. Woodley for their comments on
drafts of the manuscript.
LITERATURE CITED
Balciunas, J.K., and D.W. Burrows. 1993. The rapid
suppression of the growth of Melaleuca quinque-
nervia saplings in Australia by insects. Journal of
Aquatic Plant Management 31: 265-270.
Balciunas, J.K., D.W. Burrows, and M.F Purcell.
1994. Field and laboratory host ranges of the
Australian weevil, Oxyops vitiosa (Coleoptera:
Curculionidae), a potential biological control
agent for the paperbark tree, Melaleuca quinque-
nervia. Biological Control 4: 351—360.
1995. Insects to control Melaleuca II: pros-
pects for additionals agents from Australia.
Aquatics 17 (2):16—21.
Balciunas, J.K., and T.D. Center. 1991.
control of Melaleuca quinquenervia: prospects
and conflicts, pp. 1—2. Jn Center, T.D. et al., eds.,
Proceedings, Symposium on Exotic Pest Plants,
University of Miami, Florida, 2—4 November
1988. United States Department of the Interior,
National Park Service, Denver, Colorado.
Barlow, B.A. 1988. Patterns of differentiation in trop-
ical species of Melaleuca L. (Myrtaceae). Pro-
ceedings of the Ecological Society of Australia
15: 239-247.
Blake, S.T. 1968. A revision of Melaleuca leucaden-
dron and its allies (Myrtaceae). Contributions
from the Queensland Herbarium No. |. Queens-
land Herbarium, Department of Primary Indus-
tries, Brisbane. 114 pp.
Bodkin, EF 1991. Encyclopaedia Botanica: The Essen-
tial Reference Guide to Native and Exotic Plants
in Australia. Cornstalk Publishing, North Ryde,
Australia 1083 pp.
Bodle, M.J., A.P. Ferriter, and D.D. Thayer. 1994. The
biology, distribution, and ecological consequences
of Melaleuca quinquenervia in the Everglades, pp.
341-355. In Davis S.M. and J.C.Cogden, eds. Ey-
Biological
>)
1°)
£
erglades The Ecosystem and Its Restoration. St.
Lucie Press.
Florida Conservation Foundation. 1993. Guide to
Florida Environmental Issues and Information.
Winter Park, Florida. 364 pp.
Gagné, R.J. 1989a. The Plant-Feeding Gall Midges of
North America. Cornell University Press, Ithaca,
New York. xi and 356 pp., 4 pls.
. 1989b. Family Cecidomyiidae, pp. 152-163.
In Evenhuis, N.L., ed. A Catalog of the Diptera
of the Australasian and Oceanian Regions. Bishop
Museum Press and E. J. Brill, Honolulu, Hawaii.
1994. The Gall Midges of the Neotropical
Region. Cornell University Press, Ithaca, New
York. xv and 352 pp.
. 1995. Revision of tetranychid (Acarina) mite
predators of the genus Feltiella (Diptera: Ceci-
domyiidae). Annals of the Entomological Society
of America 88: 16—30.
Gagné, R. J., and J. Marohasy. 1994. The gall midges
(Diptera: Cecidomyiidae) of Acacia spp. (Mimo-
saceae) in Kenya. Insecta Mundi 7: 77-124.
Harris, K. M. 1966. Gall midge genera of economic
importance (Diptera: Cecidomyiidae). Part I: In-
troduction and subfamily Cecidomylinae; super-
tribe Cecidomyiidi. Transactions of the Royal En-
tomological Society of London 118: 313-358.
1979. Descriptions and host ranges of the
sorghum midge,
Contarinia sorghicola (Coquillett) (Diptera: Cecido-
myiidae), and of eleven new species of Contarinia
reared from Gramineae and Cyperaceae in Aus-
tralia. Bulletin of Entomological Research 69:
161-182.
Holliday, I. 1989. A Field Guide to Melaleucas. Ham-
lyn, Australia. 254 pp.
Kolesik, P. 1995a. Skusemyia allocasuarinae, a new
genus and species of Cecidomyiidae (Diptera)
damaging lateral branch buds of drooping sheoak,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Allocasuarina verticillata in Australia. Transac-
tions of the Royal Society of South Australia 119:
41-46.
. 1995b. A new species of Eocincticornia Felt
(Diptera: Cecidomyiidae) on Eucalyptus fascicu-
losa in South Australia. Journal of the Australian
Entomological Society 34: 147-152.
McAlpine, J.E 1981. 2. Morphology and terminolo-
gy—adults, pp. 9-63. In McAlpine, J.E et al., eds.
Manual of Nearctic Diptera, Vol. 1., Research
Branch, Agriculture Canada, Monograph 27.
Rockwood, D.L., and TE Geary. 1991. Growth of 19
exotic and two native tree species on organic soils
in southern Florida, pp. 283-302. Jn Center, T.D.
et al., eds., Proceedings, Symposium on Exotic
Pest Plants, University of Miami, Florida, 2—4 No-
vember 1988. United States Department of the In-
terior, National Park Service, Denver, Colorado.
Schiner, I.R. 1868. Diptera. [Art 1], 388 pp., 4 pls. Jn
Reise der 6sterreichischen Fregatte Novara. Zool-
ogie, Vol. 2, Abt. 1, [Sect.] B. Wien.
Schmitz: “DiC; Biv-" Nelson? I.E. Nall® andeieb:
Schardt. 1991. Exotic aquatic plants in Florida: a
historical perspective and review of the present
aquatic plant regulation program, pp. 303-326. In
Center, T.D. et al., eds., Proceedings, Symposium
on Exotic Pest Plants, University of Miami, Flor-
ida, 2-4 November 1988. United States Depart-
ment of the Interior, National Park Service, Den-
ver, Colorado.
Skuse, EA.A. 1888. Diptera of Australia. Proceedings
of the Linnean Society of New South Wales (2)
3: 17-145, pls. 2-3.
1890. Diptera of Australia. Nematocera.—
Supplement I. Proceedings of the Linnean Society
of New South Wales (2) 5: 373-413.
Yukawa, J., and S. Haitsuka. 1994. A new cecido-
myiid successor (Diptera) inhabiting empty midge
galls. Japanese Journal of Entomology 62: 709—
718.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 335-337
A NEW GENUS AND SPECIES BELONGING TO THE SUBFAMILY
STERICTIPHORINAE (HYMENOPTERA: SYMPHYTA: ARGIDAE)
FROM JAPAN
ICHIJI TOGASHI
1-chome, Honmachi, Tsurugi-machi, Ishikawa Prefecture, 920-21, Japan.
Abstract.—Aproceroides sakuraii, New Genus, New Species, is described from Japan.
The genus is separated from related genera of Sterictiphorinae, and a key is given to the
genera of Sterictiphorinae from Japan.
Key Words:
In this paper, I describe a new genus and
new species of the argid subfamily Steric-
tiphorinae from Japan. A revised key to the
Japanese genera of the subfamily will sep-
arate this new genus from other genera. The
holotype is deposited in the collection of
the National Science Museum (Natural His-
tory), Tokyo.
Aproceroides Togashi, new genus
Robust. Head from above transverse
(Fig. 2); eyes slightly converging below
(Fig. 1); space between antennal sockets
narrow, with a longitudinal ridge in middle
(Fig. 1); malar space broad (Fig. 1); clypeus
nearly truncate (Fig. |); labrum small; cen-
chri large. Forewing: apical portion of costa
clearly swollen (Fig. 3); first cubital cross
vein absent (Fig. 3); third cubital cross vein
curved; basalis joining subcosta at a dis-
tance removed from begining of cubitus
nearly equal to length of first cubital cross
vein; anal cell of forewing broadly con-
stricted in middle with small basal anal cell.
Hindwing: analis short, not connected with
brachius, with open anal cell (Fig. 4). Legs:
tarsal claw simple.
Type species: Aproceroides sakuraii sp.
nov.
This new genus is very closely allied to
Symphyta, Sterictiphorinae, Aproceroides
the genus Sterictiphora Billberg, but it is
easily distinguished from the latter by the
presence of the short anatis in hindwing and
open anal cell (in Sterictiphora, the analis
of the hindwing is complete and connected
with the brachius, with a closed anal cell),
and by the swollen apical portion of the
costa of forewing (in Sterictiphora, the api-
cal portion of costa is not swollen).
From the genus Aproceros Malaise, it is
at once distinguished by the broadly con-
tracted anal cell of forewing (in Aproceros,
the anal cell of forewing is petiolate without
a basal anal cell).
From the genus Yasumatsua Togashi, it
is easily separable by the simple tarsal claw
(in Yasumatsua, the tarsal claw has a basal
lobe).
Aproceroides sakuraii Togashi,
new species
(Japanese name: Sakurai-kuwagata-
habachi)
(Figs. 1—7)
Male.—Length 4 mm. Body black, with
following parts yellow: latero-posterior por-
tion of pronotum and tegula, and following
parts dirty yellow: labrum, maxillary and
labial palpi, and propodeum. Wings hya-
line, basal %4 of costa of forewing pale yel-
336
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-7.
low, apical %4 of costa, stigma and other
veins brownish black to black. Legs yellow
but all coxae and trochanters black, basal
half of all femora brownish black, apical
portion of hind tibiae and all tarsi brownish
black.
Head: 0OL:POL:0CL = 0.9:1.0:0.7; post-
ocellar area convex; postocellar furrow dis-
tinct but interrupted in middle (Fig. 2); lat-
eral furrows slightly depressed; circumo-
cellar furrow distinct (Fig. 2); frontal area
gently convex; median fovea ellipsoidal in
form, deep, with a median longitudinal fur-
row (Fig. 1); lateral foveae distinct; supra-
clypeal area convex; clypeus narrow; malar
space broad, nearly as long as % diameter
of front ocellus (ratio about 1.0:2.0).
Aproceroides sakuraii. 1, Head, front view. 2, Head, dorsal view. 3, Forewing. 4, Hindwing. 5,
Inner front tibial spur, lateral view. 6, Hind femur and tibia, lateral view. 7, Hind tarsus, lateral view. Scale: 0.5
mm for Figs. 1, 2, and 7; 0.1 mm for Fig. 5; 1.0 mm for Figs. 3, 4, and 6.
Thorax: Normal. Wings: venation of
forewing as in Fig. 3; hindwing with 2 mid-
dle cells; analis of hindwing short, not con-
nected with brachius, with open anal cell
(Fig. 4). Legs: hind basitarsus as long as
following 3 segments combined (ratio about
1.0:1.0); inner hind tibial spur broad (Fig.
7), and slightly shorter than % length of
hind basitarsus (ratio about 1.0:2.3); inner
front tibial spur longer than apical width of
front tibia (ratio about 1.0:0.5) (Fig. 5).
Abdomen: Normal.
Punctures.—Head and thorax nearly im-
punctate, shining. Abdominal tergites near-
ly impunctate.
Female.—Unknown.
Food plant.—Unknown.
VOLUME 99, NUMBER 2
Habitat.—Japan (Honshu).
Holotype: Male; April 17, 1994; Ameya,
Aizuwakamatsu City, Fukushima Pref., S.
Sakurai leg.
KEY TO THE GENERA OF JAPANESE
STERICTIPHORINAE
1. Tarsal claws simple
— Tarsal claws with a basal lobe at base; anal cell
of forewing petiolate; hindwing with a closed
amr al Cella eerie: cuts och access ols Yasumatsua Togashi
2. Anal cell of forewing petiolate without a basal
anal cell; basalis of forewing joins subcosta at
a point much closer to the origin of cubitus
than length of first cubital cross vein
Spe Pees a hele Ae eiiies oe Ts Tak Aproceros Malaise
— Anal cell of forewing broadly constricted in the
middle, with basal anal cell; basalis of fore-
wing joins subcosta at a distance removed from
the begining of cubitus that is a little longer
than the length of first cubital cross vein (Fig.
SO) RPeSRy Li tth it eye ey th POA os ly be 3
3. Hindwing with a closed anal cell (analis vein
ie)
ies)
~
connected to brachius); apex of costa of fore-
wing not swollen Sterictiphora Billberg
— Hindwing with open anal cell (Fig. 4) (analis
vein not connected to brachius); apical portion
of costa of forewing swollen (Fig. 3)
sp b16 6 £5 68.0 D\s GN! CREE Aproceroides, n. gen.
ACKNOWLEDGMENTS
I cordially thank Dr. D. R. Smith, U. S.
Department of Agriculture, Washington, D.
C., for review of the manuscript. I am in-
debted to Mr. S. Sakurai, Niitsu City, Nu-
gata Prefecture, for sending me the valuable
specimen.
LITERATURE CITED
Togashi, I. 1968. Japanese sawflies of the subfamily
Sterictiphorinae (Hymenoptera, Symphyta). Kon-
ty 36: 285-293.
1970. New genus and species of the Steric-
tiphorinae (Hym., Argidae) from Japan. Mushi 44:
49-53.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 338-347
THE GENUS CARPOMYA COSTA (DIPTERA: TEPHRITIDAE):
NEW SYNONYMY, DESCRIPTION OF FIRST AMERICAN SPECIES,
AND PHYLOGENETIC ANALYSIS
ALLEN L. NORRBOM
Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart-
ment of Agriculture % National Museum of Natural History, MRC 168, Washington, DC
20560, U.S.A.
Abstract.—Carpomya tica, new species, from Costa Rica is described and reported as
the first species of Carpomya from the Americas. Phylogenetic relationships within Car-
pomya are analyzed. Myiopardalis Bezzi and Goniglossum Rondani are recognized as
subjective junior synonyms of Carpomya. Carpomya wiedemanni, n. comb., and C. par-
dalina (originally described in Carpomya) are transferred to Carpomya from Goniglossum
and Myiopardalis, respectively, and Spilographa caucasica Bigot is removed from the
genus. A lectotype is designated for C. pardalina, and a key to the species of Carpomya
is provided.
Resumen.—Se describe Carpomya tica, especie nueva, de Costa Rica, la primera es-
pecie de Carpomya de las Américas. Se analizan las relaciones filogeneticas dentro de
Carpomya. Myiopardalis Bezzi y Goniglossum Rondani son sinonimos subjectivos nuevos
de Carpomya. Se transfieren C. pardalina (descrita originalmente en Carpomya) y Car-
pomya wiedemanni, n. comb., a Carpomya de Myiopardalis y Goniglossum, respectiva-
mente, y se remueve Spilographa caucasica Bigot de este género. Se designa un lectotipo
para C. pardalina, y se provee una clave a las especies de Carpomya.
Key Words: Fruit flies, Tephritidae, Carpomya, Myiopardalis, Goniglossum
The genus Carpomya Costa (1854) pre- METHODS
viously included four species known from
the southern Palearctic, northeastern Afro-
tropical, and western Oriental Regions, al-
though one of these species, C. caucasica
(Bigot) does not seem to belong in this ge-
I follow the morphological terminology
of McAlpine (1981), except as noted by
Norrbom and Kim (1988). Terminology for
the wing pattern follows Foote (1981, see
Fig. 71). Taxonomically useful characters
nus. The monotypic genera Myiopardalis
Bezzi (1910) and Goniglossum Rondani
(1856), here considered new subjective ju-
nior synonyms of Carpomya, included one
southern Palearctic/western Oriental species
and one Palearctic: species, respectively.
This paper reports the discovery of the first
American species of Carpomya, increasing
the number of species of this genus to six,
and substantially extending its distribution.
within Carpomya are listed in Table 1 and
the distributions of their states are shown in
Table 2, but only the nine characters with
an asterisk were used in the phylogenetic
analysis; the others are autapomorphies.
The Hennig86 program was used for phy-
logenetic analysis, with the Rhagoletis ta-
bellaria group as the outgroup for assigning
character polarities. Transformation series
and the polarity of some characters are fur-
VOLUME 99, NUMBER 2 339
Table 1. Characters taxonomically useful for species of Carpomya. Only characters with an * were used in
phylogenetic analysis.
ao:
. First flagellomere—O) with dorsoapical point; 1) rounded dorsoapically.
. Mesonotal color and microtrichia pattern—O) unicolorous except white areas, or if partially brown without
*6.
Tf
ao:
A),
10.
ilile
12:
== I13}-
14.
115).
16.
eali//.
. Head elongate, gena especially produced and angulate anteriorly, and proboscis long geniculate—O) no;
1) yes.
. Facial carina—O) moderately to strongly produced dorsally, gradually becoming broader and less produced
ventrally; 1) weakly produced, gradually becoming broader ventrally; 2) strongly produced dorsally, rather
abruptly becoming very broad and weakly produced near midlength; 3) extremely produced and moderately
broad throughout, carinate on both sides. States 1, 2, and 3 are here considered independently derived
autapomorphies.
Ocellar setae—0O) well developed; 1) minute, length less than width of ocellar tubercle.
microtrichia or with microtrichia unicolorous; 1) with pattern of dark brown or black marks (similar to
Fig. 1), some without microtrichia, others with dense black microtrichia.
Scutellar color pattern—O) brown basally, white apically; 1) with medial brown spot extended to apex or
with smaller isolated medial brown spot and apical brown spot(s); 2) without medial brown spot, pale area
m-shaped, with medial pale arm; 3) entirely yellow. The transformation series of this character is uncertain
and it was coded unordered in the Hennig86 analysis. Carpomya tica, pardalina and wiedemanni were all
coded 1, although there are some differences in their scutellar patterns: C. tica has a large medial brown
spot and U-shaped yellow area which includes the apical setae (Fig. 1); C. pardalina has a smaller medial
brown spot, isolated by a larger pale area from an apical brown spot or spots on which the apical setae
are located; and C. wiedemanni is variable, either like C. pardalina, or with the medial brown spot large
and including the apical setae (Freidberg & Kugler 1989: 191).
Scutellum shape—O) relatively triangular in dorsal view, disc flat to slightly convex; 1) almost semicircular
in dorsal view, disc slightly to moderately convex (a little less convex in incompleta than in schineri and
vesuviana); 2) almost semicircular in dorsal view, disc very strongly convex.
Katepisternum—Q) unicolorous brown (R. tabellaria group) or yellow (C. tica); 1) yellow or orange, with
dorsal margin white (contrast often weak in C. incompleta and pardalina); 2) with large brown or black
area, dorsal margin white. State 0 appears to be plesiomorphic, although it should be noted that states |
and 2 occur in Zonosemata.
Cell r, with spur vein—O) with at most a dark spot or crease within the subapical band; 1) with a distinct
spur vein originating from vein R,,, in the subapical band.
Crossvein r-m—Q) near midpoint between bm-cu and dm-cu, within or at apical margin of discal band; 1)
at 0.73 distance from bm-cu to dm-cu, touching proximal margin of subapical band.
Subbasal and discal bands—O) not connected or connected posteriorly; 1) connected in cells r, and br.
Anterior apical wing band—Q) present; 1) absent.
Epandrium—0) mostly brown; 1) mostly yellow with small dorsomedial brown spot. In other mostly yellow
Carpomyini the epandrium is mostly brown or all yellow or has paired dorsolateral spots, although in
Zonosemata it varies from states 0 to 1, sometimes intraspecifically.
Aculeus tip—QO) nonserrate; |) serrate.
Spermatheca number—0O) 3; 1) 2. The number of spermathecae varies from 2—3 in the R. tabellaria group
and other Rhagoletis and Carpomyini (Bush 1966, Norrbom 1994b), but 3 is the plesiomorphic state for
Tephritidae, so I tentatively coded that number as plesiomorphic within Carpomya and assigned that state
to the outgroup in the Hennig86 analysis.
Spermatheca shape—O) single chambered; 1) double chambered.
Host plant—0O) Rosa spp. (Rosaceae); 1) Ziziphus spp. (Rhamnaceae); 2) Cucurbitaceae (C. paradalina
breeds in Cucumis, Citrullus, and Ecballium spp., and C. wiedemanni in Bryonia spp.). The transformation
series is uncertain; states 1 and 2 are probably apomorphic, as C. pardalina and C. wiedemanni are the
only two species of Carpomyini known to breed in Cucurbitaceae, and C. incompleta and C. vesuviana
are the only Carpomyini known from Rhamnaceae (Freidberg & Kugler 1989, White & Elson-Harris 1992),
but these states may be independently derived from state 0 or some other state (a wide range of host
families are attacked by Carpomyini). The hosts of C. tica are unknown. Coded unordered in the Hennig86
analysis.
340 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 2. Character state distributions among species of Carpomya.
ee ee ee e———————————
Characters
1 1 1 1 1 1 1
Taxa 2 3 4 5 6 7 8 9 0 I 2 3 4 5 6 7
Se ee —————————————— eee
outgroup 0 0/1 0 —. OOo OF Ok PAO a Oe. OF Ora Og te O/1 0) ?
tica 0) i 0) 10) 1 2) 0) 0 1 0) 0) 10) 2 uy ? ey
pardalina O ) O O 1 | O 1 1 0) 0) 0) 1 O 0) l 2
wiedemanni | 3 0) 1 1 1 0) 2 ] 0) | 0) 1 ] 0) 10) 2
schineri 0) 0) 0) 0) 1 2 1 2 0) 0) 0) 0) 1 0) 1 0) 0
incompleta O O ] 0) O 3 i 1 0) 0) 0) l 1 0) | 0) 1
vesuviana O O | 0) 1 2 1 1 0) 0) 0) 0) 1 0) 1 0) 1
ther discussed in Table 1. Note that char- band of wing present. Ocellar seta well devel-
acters 6 and 17 were treated as unordered. Oped OF TAIL: 12 42g: ncleg i ora eae =
TAXONOMY
KEY TO THE SPECIES OF CARPOMYA
1. Vein R,,, with a distinct, anteriorly directed
spur vein in the subapical band. Three sper-
mathecae. Host plants Cucurbitaceae. ....... 2
— Vein R,,, without a spur vein, cell r, with at
most a dark spot or crease within the subapical
band. Two spermathecae (unknown for C.
tica). Host plants not Cucurbitaceae (unknown
for C. tica).
2. Head elongate, gena especially produced and
angulate anteriorly. Proboscis long geniculate.
Facial carina strongly produced throughout and
only slightly wider ventrally than dorsally. Ka-
tepisternum with a large dark brown spot me-
dially. Subbasal and discal bands connected in
cells r, and br. Aculeus tip serrate. Europe, Is-
LAGI S.. (5 eA eee eros wiedemanni (Meigen)
— Head not elongate, gena rounded anteriorly.
Proboscis short, capitate. Facial carina strongly
produced dorsally, but ventral half very broad
and weakly produced. Katepisternum yellow or
orange, except for whitish dorsal margin. Sub-
basal and discal bands not connected. Aculeus
tip nonserrate. Caucasus, Turkey, Cyprus and
Egypt to western India. ....... pardalina Bigot
3. Crossvein r-m near midpoint between bm-cu
and dm-cu, within or at apical margin of discal
band. Scutellum dark brown with m-shaped
whitish area, including a medial pale mark, or
enimelyayellow- Old a VWioOrldss aii) iene nee 4
— Crossvein r-m at about 4 distance from bm-cu
to dm-cu, touching proximal margin of sub-
apical band (Fig. 2). Scutellum dark brown
with U-shaped yellowish mark (Fig. 1), with-
out medial pale mark. Host plants unknown.
@ostagRicaly a. istire sas oom eters tica, N. sp.
4. Scutum and scutellum with large dark brown
or black spots or markings. Anterior apical
— Scutum and scutellum without brown or black
spots or markings. Anterior apical band of
wing absent. Ocellar seta minute, length less
than width of ocellar tubercle. Host plants Ziz-
iphus spp. Southern Europe, Middle East,
Egypt, Sudan, Ethiopia. ... . incompleta (Becker)
5. Ocellar seta well developed. Katepisternum
with at least a large dark brown spot medially,
usually mostly dark brown except whitish dor-
sal margin. Host plants Rosa spp. Central Eu-
rope to Kazakstan and Israel. .. . schineri (Loew)
— Ocellar seta minute, length less than width of
ocellar tubercle. Katepisternum yellow or or-
ange, except for whitish dorsal margin. Host
plants Ziziphus spp. Italy, Bosnia, Caucasus,
Central Asia, Pakistan, India, Thailand.
i sip yicia Fonsi heen ar OTe gene tae cue vesuviana Costa
Carpomya pardalina Bigot
Carpomyia pardalina Bigot 1891: 51.
Myiopardalis pardalina: Bezzi 1910: 11;
Freidberg and Kugler 1989: 194; White
and Elson-Harris 1992: 349. See latter
two publications for additional refer-
ences.
Carpomyia (Myiopardalis) pardalina:
Zaitzev 1919: 66; Rohdendorf 1939: 8;
Kandybina 1965: 668 [larva].
Carpomyia (Myiopardalis) caucasica Zait-
zev 1919: 64; Stackelberg 1928: 281
[synonymy]; Zaitzev 1947: 6 [synony-
my].
Bigot described C. pardalina from “‘plu-
sieurs’’ (several) specimens of both sexes
from ‘‘Beloutchistan’’ (Baluchistan, an area
extending from southeastern Iran to western
VOLUME 99, NUMBER 2
Pakistan). In the Bigot Collection, now at
the University Museum, Oxford, there are
three specimens (2 ¢ 1 @) on separate pins,
one of each sex also with a larva pinned
below the adult specimen. The pin of the
female, here designated as lectotype, has a
thin paper label of the style that Ackland
and Taylor (1972, Fig. 6) found mainly on
specimens with Bigot manuscript names.
The lectotype’s label has the following data
in Bigot’s writing: “‘Carpomyia pardalina
[the second half of the specific name writ-
ten over some other letters] 2 \n. sp. Inedit.
\ qui[rest of word illegible] Octobre 1890.
J. Bigot \ Beloutchistan \ attaque les mel-
ons.”’ There is a fourth pin with a similar
label except for a “‘d”’ symbol instead of
*“®.”’ It has only some debris, perhaps the
remains of a third syntype male. The lec-
totype is pinned through its abdomen and
has slightly shriveled eyes, but otherwise is
in good condition. All of these specimens
fit Bigot’s description and the traditional in-
terpretation of this species.
Carpomyia caucasica Zaitzev (1919)
was described from male and female spec-
imens from ‘East Transcausia, Dzhevan-
shir, Areshsk territory, Elisavetn region and
North Mugan.”’ Elisavetn is probably the
locality in Azerbaijan that has also been
known as Gandzhe, Kirovabad, and Yeli-
savetpol (A. Konstantinov, personal com-
munication). This name has long been con-
sidered a synonym of C. pardalina by Rus-
sian workers (Stackelberg 1928, Zaitzev
1947, Kandybina 1965), but has been little
noticed by western authors. It was omitted
from the Palearctic Diptera catalog (Foote
1984). I examined a pair of specimens of
C. pardalina in the Zoological Institute, St.
Petersburg with the following label data:
*A3. o. 3. CT.”” [= Azerbaijan?] and hand-
written localities that I cannot decipher; the
male with the date *‘19/viii 1928”, and the
female with the date ‘*3/vili 1927.” The fe-
male also has a determination label with
*“Myiopardalis caucasica m., Zaitzev det.”
These specimens cannot be types because
their collection dates are subsequent to the
341
Fig, 1:
Carpomya tica, thorax, dorsal view.
date of description, but their identity helps
to confirm the synonymy of this name.
As explained in the Relationships sec-
tion, I follow Zaitzev (1919, 1947), Roh-
dendorf (1939) and Kandybina (1965) in
including pardalina in Carpomya, rather
than in the monotypic genus Myiopardalis,
as it has been treated by the majority of
authors.
Carpomya tica Norrbom, new species
(Figs. 1—4)
Holotype—d (at USNM, for eventual
deposit in Instituto Nacional de Biodiver-
sidad, Heredia, Costa Rica), COSTA RICA:
San José: Zurqui de Moravia, 10°03’N
84°01'W, 1600 m, Malaise trap, V.1992, P.
Hanson.
Diagnosis.—Carpomya tica differs from
all other species of Carpomya and most
other Carpomyina in the distal location of
crossvein r-m (Fig. 2), which is at the apical
¥%4 of cell dm versus near the middle of that
cell in other Carpomyina except Crypto-
dacus spp. and Haywardina cuculi (Hen-
del), which differ from Carpomya in having
342 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 2.
Carpomya tica, right wing.
a white medial scutal stripe or spot. C. tica
also differs from other species of Carpo-
myina, including most of the other species
of Carpomya, by the strongly swollen
shape of its scutellum. Only C. schineri and
C. vesuviana approach C. tica in scutellum
shape, but neither has this structure as stout
as in C. tica, and they further differ in the
shape of the pale area of their scutellar col-
or patterns, which is somewhat m-shaped,
with a medial pale mark (see Freidberg and
Kugler 1989, Fig. 188), rather than U-shaped
as in C. tica (Fig. 1; the yellow areas are
continuous on the extreme apex of the scu-
tellum, visible in posterior view). In the key
to Neotropical genera of Tephritidae in
Foote (1980), C. tica will key to Cecido-
chares, which is not closely related (it be-
longs to the subfamily Tephritinae). Species
of the latter genus differ as follows: body
mostly dark brown, including all of the scu-
tum and scutellum; at least some scutal
setulae swollen; and outer and inner surstyli
short.
Description.—Setae black. Body pre-
dominantly yellow. Head: Yellow. 3 fron-
tal, 2 orbital setae, all large. Ocellar seta
large (length about equal to width of frons).
Facial carina weak. First flagellomere about
2 times as long as wide (lateral view), with
small dorsoapical point. Proboscis short,
capitate. Thorax: Mostly yellow. Postpro-
notal lobe, dorsal margin of anepisternum,
and posterior third of notopleuron white.
Mesonotum (Fig. 1) with following dark
brown markings: medial spot from level of
dorsocentral seta to level of acrostichal seta,
extended laterally slightly beyond acrosti-
chal seta; spot on anterior two-thirds of no-
topleuron, extended mesally on scutum to
level of presutural supra-alar seta but not
including it; broad spot along transverse su-
ture; spot from wing base to slightly mesal
to level of intra-alar seta and from postsu-
tural supra-alar seta almost to post-alar and
intra-alar setae; spot posterior and lateral to
intra-alar seta; and scutellum, except for
narrow, U-shaped yellow mark that in-
cludes apical and basal setae. Pleuron with-
out dark brown markings. Subscutellum en-
tirely dark brown. Mediotergite yellow lat-
erally, broadly dark brown medially, more
broadly so dorsally. Scutellum strongly
swollen. Chaetotaxy as in Rhagoletis and
Carpomya. Dorsocentral seta slightly ante-
rior to level of postsutural supra-alar seta.
Scapular setae white. Mesonotal setulae
slender; those on scutum yellow or black,
occurring in patches. Scutellum with yellow
setulae, present only on yellow areas.
Mesonotum mostly moderately densely
pale microtrichose, except for dark brown
part of notopleuron, dark brown spot along
transverse suture, lateral half of brown spot
VOLUME 99, NUMBER 2
0.1 mm
Figs. 3—4.
Epandrium, surstyli and cercus, lateral view.
between postsutural supra-alar and intra-
alar setae, and spot posterior and lateral to
intra-alar seta, which are densely black mi-
crotrichose, and the following nonmicrotri-
chose areas: dark brown part of scutum
mesal to notopleural dark area; middle of
medial scutal brown spot; mesal half of
brown spot between postsutural supra-alar
and intra-alar setae; and scutellum except
yellow areas and base and apex of lateral
brown area. Subscutellum and upper half of
brown area of mediotergite densely black-
ish microtrichose; yellow parts of medi-
otergite moderately pale microtrichose;
ventral part of brown area bare. Legs: Yel-
low, except hind tibia with basal posterior
and subapical anterior and posterior brown
spots. Wing (Fig. 2): With subbasal, discal,
preapical and anterior apical bands. Acces-
sory costal band and posterior apical bands
absent. Discal and subapical bands parallel,
perpendicular to long axis of wing; each
broad, but not connected. Anterior apical
band only narrowly connected to subapical
3
£
WwW
Carpomya tica, male genitalia. 3, Epandrium and surstyli, posterior view (cerci not shown). 4,
band in cell r,, and separated from costa by
hyaline marginal spots in cells r, and cell
r>,3. Cell r, without spur vein from vein
R,,; in subapical band. Crossvein r-m at
0.73 distance from bm-cu to dm-cu (just
touching proximal margin of subapical
band). Cell bcu with apical lobe 0.5 times
as long as maximal width of cell. Abdomen:
Mostly yellow, each tergite (through tergite
5) with unpaired medial and paired subla-
teral brown spots, small on syntergite 1+2.
Male terminalia (Figs. 3—4): Epandrium
brown, surstyli yellow. Outer surstylus
slender, with distinct mesal lobe; part apical
to mesal lobe with relatively few small
setulae.
Spilographa caucasica Bigot
Hendel (1927) placed this species in Car-
pomya, but probably based on confusion
with Carpomyia caucasica Zaitzev (= C.
pardalina). Without examination of the ho-
lotype it is difficult to recognize S. caucas-
ica. Bigot’s description is vague and there
344
are no illustrations. But it is very doubtful
that S. caucasica belongs in Carpomya, as
no species of that genus has a longitudinal
band in addition to four transverse bands on
the wing. Bigot’s statement “‘quatre bandes
roussatres, peu distinctes, sises a la partie
postérieure du thorax”? (4 reddish bands,
not very distinct, situated at the posterior
part of the thorax), also seems questionable
for a species of Carpomya, which except
for C. incompleta (which has only 3 wing
bands and therefore cannot be S. caucasi-
ca), have distinct dark brown spots or larger
marks on the mesonotum. I cannot recog-
nize S. caucasica, and treat it here as an
unplaced species of Trypetinae.
Bigot (1880: 154, bottom of page) stated
clearly that the species he described in this
paper were in his collection, but Adrian
Pont (personal communication) was unable
to locate the single female (therefore holo-
type) of S. caucasica in the Bigot Collec-
tion at the University Museum, Oxford. He
remarked that it is possibly there under
‘““some other name that Bigot changed to
caucasica with publication. He was prone
to doing this, and it causes endless confu-
sion.”
RELATIONSHIPS AND CLASSIFICATION
The Carpomyina was proposed as a sub-
tribe of Trypetini by Norrbom (1989), but
is currently considered a subtribe of Car-
pomyini within the Trypetinae (Korneyev
1996). The desclerotized area at the apex of
female syntergosternite 7 and the presence
of stomal guards, usually distinctly sclero-
tized, in the larva are probable synapomor-
phies for the subtribe (Norrbom 1989, Car-
roll 1992). The shape of the male surstyli
(inner and outer surstyli elongate, the latter
with a long, apically directed posterior lobe
and a short, mesally directed anterior lobe)
is probably another synapomorphy of the
Carpomyina or possibly for the Carpomyini
(Korneyev 1996). Some reversal (reduction
of the posterior lobe of the outer surstylus
in a few taxa) must have occurred if this
character was in the groundplan of the Car-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
pomyina, but other character state distri-
butions support that hypothesis (Norrbom
1989, 1994b).
Carpomya, as recognized here, includes
Six species: C. incompleta (Becker), C. par-
dalina Bigot, C. schineri (Loew), C. vesu-
viana Costa, C. wiedemanni (Meigen), NEW
COMBINATION and C. fica, n. sp. Carpomya
tica belongs in the Carpomyina, or at least
to the Carpomyini, based on the shape of
its male genitalia; its surstyli have the typ-
ical carpomyine shape (Figs. 3—4). With
most of the other species of Carpomya, C.
tica shares a distinctive, apomorphic color
pattern of the cuticle and microtrichia of the
thorax, which is mostly yellow with dark
brown to black marks on the scutum, scu-
tellum, and subscutellum (Fig. 1; also see
Freidberg and Kugler 1989, Figs. 188, 202,
209). The microtrichia are pale or gray, ex-
cept on some of the dark brown areas,
which are bare or have dense, blackish mi-
crotrichia. Except for the subscutellar
marks, this pattern has been lost in C. in-
completa, which shares several synapomor-
phies with C. schineri and vesuviana (see
Fig. 5), indicating that it does belong in
Carpomya (J. Jenkins, personal communi-
cation, has discovered a male genitalic syn-
apomorphy that further supports this clade).
This color pattern is unique within the Car-
pomyina; in other genera, the thorax is dark
brown or black (except, in many species,
for the whitish postpronotal lobe, dorsal
anepisternal stripe, apical part of scutellum,
and sometimes a medial scutal spot or stripe)
or predominantly yellow to orange, or if
partly or mostly dark brown (e.g., Zonose-
mata or Rhagoletis suavis species group),
the pattern is much different than in Car-
pomya. Some species of Ceratitidina in the
Dacini do have thoracic color patterns sim-
ilar to Carpomya, thus convergent evolu-
tion of such a pattern in C. tica and the Old
World species of Carpomya is possible, but
I have discovered no apomorphies shared
by C. tica and any other species of Carpo-
myina (or other Trypetinae) that would con-
tradict the hypothesis that Carpomya, as de-
VOLUME 99, NUMBER 2
=(5,/6.. 4)
Euca
‘Cia pardalina
wiedemanni
al peead ines AL 4/ 2) a (8.2) =
(Sec 3) (8.2)=—— schineri
Apewsstiea
Fig. 5.
Table 1.
limited here, is monophyletic. Other char-
acter state distributions indicate that the dis-
tal location of r-m in C. tica and
Cryptodacus and Haywardina cuculi is the
result of convergence (see Norrbom
1994b).
Certain characters of the wing pattern
may be additional synapomorphies of Car-
pomya, although some species now placed
in Rhagoletis (particularly the tabellaria
species group) have similar wing patterns,
so some or all of these characters could be
plesiomorphic or synapomorphies at a high-
er level within the Carpomyina. Better un-
derstanding of the polarity and evolution of
wing patterns within the Carpomyina is
necessary to determine the phylogenetic
significance of these characters, but at least
the strong similarity of the wing pattern of
C. tica with those of the other species of
Carpomya does not contradict its classifi-
cation in this genus. The similar elements
of the wing pattern include: absence of the
accessory costal band; absence of the pos-
terior apical band; the broad width of all of
the bands (reduced in C. incompleta) and
their yellow to brown color; discal and sub-
apical bands transverse and parallel; and the
at least partial separation of the anterior api-
cal band from the wing margin (Fig. 2; and
Freidberg and Kugler 1989, Pl. VIII, Figs.
4, 8, 10).
The relationship of Carpomya to other
genera of Carpomyina has not been re-
solved. Its closest relatives may be within
the possibly paraphyletic genus Rhagoletis
; retinal incompleta
"be 7) veSuviana
Hypothesized phylogenetic relationships among species of Carpomya. Character numbers refer to
(Bush 1966: 451), and I tentatively used the
R. tabellaria group, which has similar wing
patterns, as the outgroup for analysis of
character polarity.
The Hennig86 analysis of relationships
within Carpomya (based on the nine char-
acters with an asterisk in Table 1) resulted
in the single tree shown in Figure 5 (length
17 steps, consistency index 82, retention in-
dex 75). Carpomya incompleta, schineri
and vesuviana form one monophyletic sub-
group, and C. pardalina and wiedemanni
form another. These groups together appear
to be the sister group of C. tica. Despite
having several distinctive autapomorphies
(greatly elongated mouthparts and head, fa-
cial carina shape, apically rounded first fla-
gellomere, and serrate aculeus tip; see
Freidberg and Kugler 1989, Figs. 201, 204),
C. wiedemanni, the type species of Gonig-
lossum, appears to be most closely related
to C. pardalina, which is the type species
of Myiopardalis. The synonymy of Gonig-
lossum and Myiopardalis with Carpomya
(type species C. vesuviana) is a subjective
decision of ranking, but their continued rec-
ognition as monotypic genera (or the place-
ment of both species in Goniglossum)
would require the proposal of yet another
monotypic genus for C. tica. Considering
the diversity of species now classified in
Rhagoletis, I find the latter option undesir-
able. Already, Zaitzev (1919, 1947), Roh-
dendorf (1939) and Kandybina (1965,
1977) considered Myiopardalis a subgenus
of Carpomya. The latter author found little
346
difference among the larvae of the species
here included in Carpomya, except that the
oral ridges are reportedly nonserrate in C.
wiedemanni. This character (not verified by
personal examination by Kandybina in C.
wiedemanni) varies in other genera (e.g.,
Anastrepha, see White and Elson-Harris
1992), and Kandybina otherwise considered
C. wiedemanni larvae most similar to those
of C. pardalina and Rhagoletis flavigenu-
alis Hering of the tabellaria species group.
BIOGEOGRAPHY
The Carpomyina are distributed predom-
inantly in the Holarctic Region and in high-
er altitude or temperate areas of the Neo-
tropical Region. Although the Tephritidae
and most of its major clades probably orig-
inated in the Old World, the discovery of
C. tica in the Neotropics lends further sup-
port to the hypothesis that the Carpomyina
originated and diversified in the Americas,
with subsequent reinvasion of the Old
World (Norrbom 1994a). Although a thor-
ough phylogenetic analysis of the Carpo-
myina is needed to test this hypothesis of
origin, it is supported by diversity data for
the group. Except for the little known
monotypic South African genus Scleropi-
thus Munro, which appears to belong in the
Carpomyina (A. Freidberg, personal com-
munication), all of the genera of Carpo-
myina are now known from the Americas.
This includes all of the species groups of
the large, probably paraphyletic genus
Rhagoletis, except for the cerasi group.
Furthermore, all of these taxa except Car-
pomya and the alternata and tabellaria spe-
cies groups of Rhagoletis are more diverse
in the New World (Berlocher and Bush
1982, Norrbom 1994b).
ACKNOWLEDGMENTS
I am grateful to Paul Hanson for allowing
me to study the interesting Costa Rican te-
phritids taken in his Malaise traps, includ-
ing the unique holotype of C. tica. Adrian
Pont arranged the loan of the types of C.
pardalina and searched for the holotype of
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Spilographa caucasica, and V. EF Zaitzev
arranged the loan of specimens from the
Zoological Institute, St. Petersburg. Alex-
ander Konstantinov and Raymond Gagné
translated pertinent sections of Russian and
French publications. John Jenkins and Am-
non Freidberg kindly shared some unpub-
lished data (a genitalic synapomorphy con-
firming the monophyly of the vesuviana
group, and notes on the classification of
Scleropithus, respectively), as well as their
thoughts regarding the relationships and
classification of the Carpomyina. They, V.
Hernandez-Ortiz, G. J. Steck, and EF C.
Thompson reviewed earlier drafts of the
manuscript.
LITERATURE CITED
Ackland, D. M. and E. Taylor. 1972. Notes on the
Verrall-Collin Collection of Diptera in the Hope
Department of Entomology, University Museum,
Oxford. Entomologist’s Monthly Magazine 15:
12-15.
Berlocher, S. H. and G. L. Bush. 1982. An electro-
phoretic analysis of Rhagoletis (Diptera: Tephrit-
idae) phylogeny. Systematic Zoology 31: 136—
ISIS),
Bezzi, M. 1910. Restaurazione del genere Carpomyia
(Rond.) A. Costa. Bollettino del Laboratorio di
Zoologia Generale e Agraria della Rebia Scuola
d’ Agricoltura, Portici 3: 273-313.
Bigot, J. M. E 1880. Diptéres nouveaux ou peu con-
nus. 13e Partie. Quelques Diptéres de Perse et du
Caucase. Annales de la Société Entomologique de
France, Ve Série 1880: 139-154.
1891. I1Il—The Baluchistan Melon Fly.
(Carpomyia Pardalina, 3 et 2, nov. sp.). Indian
Museum Notes 2: 51.
Carroll, L. E. 1992. Systematics of fruit fly larvae
(Diptera: Tephritidae). Ph.D. dissertation, Texas A
& M University, College Station, 341 p.
Costa, A. 1854. Frammenti di entomologia napole-
tana. Annali Scientifici (Napoli) 1: 69-91.
Foote, R. H. 1980. Fruit fly genera south of the United
States (Diptera: Tephritidae). U.S. Department of
Agriculture, Technical Bulletin No. 1600, 79 pp.
. 1981. The genus Rhagoletis Loew south of
the United States (Diptera: Tephritidae). U.S. De-
partment of Agriculture, Technical Bulletin No.
1607, 75 pp.
1984. Family Tephritidae, p. 66-149. In
Soos, A. and L. Papp, eds., Catalogue of Palaearc-
tic Diptera, Vol 9, Micropezidae -Agromyzidae.
Elsevier Science Publishers, Amsterdam. 460 p.
Freidberg, A. and J. Kugler. 1989. Fauna Palaestina.
VOLUME 99, NUMBER 2
Insecta IV. Diptera: Tephritidae. Israel Academy
of Sciences and Humanities, Jerusalem, 212 pp.
Han, H.-Y. 1992. Classification of the tribe Trypetini
(Diptera: Tephritidae: Trypetinae). Ph.D. thesis,
The Pennsylvania State University, University
Park, 275 pp.
Hendel, E 1927. Trypetidae. /n Lindner, E., ed., Die
Fliegen der palaearktischen Region. Vol. 5, 221
Pp-
Kandybina, M. N. 1965. On the larvae of fruit-flies
of the genus Carpomyia A. Costa (Diptera, Try-
petidae). Enomologicheskoe Obozrenie 44: 665—
672. [In Russian, English translation in Entomo-
logical Review 44: 390-394. ]
1977. Lichinki plodovykh mukh-pestrokry-
lok (Diptera, Tephritidae) [Larvae of fruit-infest-
ing fruit flies (Diptera, Tephritidae).] Opredeliteli
Po Faune SSSR. No. 114: 1—210.
Korneyev, V. A. 1996. Reclassification of Palaearctic
Tephritidae (Diptera). Communication 3. Vestnik
Zoologii 1995(5—6): 25—48.
McAlpine, J. EF 1981. Morphology and terminology—
Adults, pp. 9-63. Jn McAlpine, J. F et al., coords.,
Manual of Nearctic Diptera, vol. 1. Agriculture
Canada, Monograph No. 27, Ottawa.
Norrbom, A. L. 1989. The status of Urophora acuti-
cornis and U. sabroskyi (Diptera: Tephritidae).
Entomological News 100: 59—66.
1994a. New genera of Tephritidae (Diptera)
from Brazil and Dominican Amber, with phylo-
genetic analysis of the tribe Ortalotrypetini. Insec-
ta Mundi 8: 1-15.
1994b. New species and phylogenetic anal-
347
ysis of Cryptodacus, Haywardina, and Rhagole-
totrypeta (Diptera: Tephritidae). Insecta Mundi 8:
37-65.
Norrbom, A. L. and K. C. Kim. (988. Revision of the
schausi group of Anastrepha Schiner (Diptera: Te-
phritidae), with a discussion of the terminology of
the female terminalia in the Tephritoidea. Annals
of the Entomological Society of America 81: 164—
7/3.
Rohdendorf, B. B. 1939. Differences between the
melon fly [Carpomyia (Myiopardalis) pardalina
Big.] and closely related species. Inf. byull. vopr.
karantina rast. 6: 8-9. [Not seen, copied from
Kandybina 1965.]
Rondani, C. 1856. Dipterologiae Italicae prodromus.
Vol 1: Genera Italica ordinis dipterorum ordinatim
disposita et distincta et in familiae et stirpes ag-
gregata. Parmae [Parma], 228 pp.
Stackelberg, A. 1928. Zur Synonymie der Melonen-
fliege Myiopardalis pardalina Big. = caucasica
Zaitz. Reports of the Bureau of Applied Ento-
mology 3: 281.
White, I. M. and M. Elson-Harris. 1992. Fruit Flies
of Economic Significance: Their Identification and
Bionomics. C.A.B. International, Wallingford, 601
PPp-
Zaitzev, EK A. 1919. The Transcaucasian melon fly—
Carpomyia (Myiopardalis) caucasica, sp. n. (Dip-
tera, Trypetidae). Zapiski nauchno-prikladnykh ot-
dieloy Tiflisskago botanicheskago sada 1: 64—66.
. 1947. The fruit fly fauna of the Caucasus and
adjacent regions (Diptera Trypetidae). Trudy
Zoologicheskogo Instituta, Akademiya Nauk,
Gruzinskoi SSR 7: 1-16.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 348-358
REVIEW OF THE BLASTOBASINAE (LEPIDOPTERA: GELECHIOIDEA:
COLEOPHORIDAE) OF THE GALAPAGOS ISLANDS
DAVID ADAMSKI AND BERNARD LANDRY
(DA) 6033 Majors Lane, Apt. #2, Columbia, MD 21045, U.S.A.; (BL) Department of
Environmental Science, Policy and Management, 201 Wellman Hall, University of Cali-
fornia, Berkeley, CA 94720, U.S.A.
Abstract.—Calosima darwini, new species, is described from the Galapagos Islands,
Ecuador, and is the first member of the genus known beyond North America. Auximobasis
normalis Meyrick, 1918 is a senior synonym of Blastobasis crotospila Meyrick, 1926,
and is redescribed and transferred to Blastobasis Zeller, 1855. A key and illustrations of
adults, including male and female genitalia, are provided for the two species of Blasto-
basinae known from the Galapagos Islands.
Key Words:
South America, Ecuador, Galapagos, Lepidoptera, Gelechioidea, Coleo-
phoridae, Blastobasinae, Calosima, Blastobasis
The Lepidoptera of the Galapagos Is-
lands are known incompletely. Although a
number of collections have been made and
reported, they have been restricted to the
larger moths and the butterflies (Beebe
1923, Butler 1877, Schaus 1923, Hayes
1975). Consequently, the microlepidoptera
remain known only partially. Eleven spe-
cies were recorded by Meyrick (1926), and
four were added by Linsley and Usinger
(1966) and Linsley (1977). Recent collec-
tions by Robert Silberglied in 1969 and
more recently by Bernard Landry in 1989
and 1992 have resulted in considerably
more specimens of microlepidoptera than
have been available previously. For exam-
ple, Landry amassed more than 3,000 spec-
imens of microlepidoptera in 5 months of
collecting on the Galapagos Islands. Studies
of this material have yielded publications
on Pterophoridae (Landry and Gielis 1992,
Landry 1993), Choreutidae (Heppner and
Landry 1994a), Heliodinidae (Heppner and
Landry 1994b), and the present paper on
Coleophoridae (Blastobasinae).
The Blastobasinae are probably one of
the most commonly collected groups of Ge-
lechioidea in the Americas. Yet this sub-
family may be one of the least known to
science. Generally, species are drab with
few, if any, diagnostic wing color patterns,
making identifications difficult unless the
genitalia are examined.
Since Meyrick (1894) the Blastobasinae
have been considered to be a monophyletic
group. Recent studies (Adamski and Brown
1989, Hodges, in press) have corroborated
this notion and have rigorously established
relationships of the Blastobasinae within
Gelechioidea. In this work the Blastobasi-
dae (sensu Adamski and Brown, 1989) are
treated as a subfamily within the Coleo-
phoridae.
MATERIALS AND METHODS
Collecting methods and information on
the islands visited are found in Landry and
Gielis (1992) and Landry (1993). Speci-
mens were prepared (pinned and mounted)
VOLUME 99, NUMBER 2
in the field as demonstrated by Landry and
Landry (1994).
The Methuen Handbook of Colour (Kor-
nerup and Wanscher, 1978) is used as a col-
or standard for the description of the adult
vestiture. More than 150 specimens of Blas-
tobasinae were examined, including type
specimens. Genitalia were dissected as de-
scribed following Clarke (1941), except
mercurochrome and chlorazol black were
used as stains. Pinned specimens and gen-
ital preparations were examined with dis-
secting and compound microscopes. Wing
measurements were made using a calibrated
ocular micrometer.
RESULTS
KEY TO THE BLASTOBASINAE
(COLEOPHORIDAE) OF THE GALAPAGOS
ISLANDS
iba IMIBIS kee euatokancnc ene com pchc orator once as ie oe 2
— Female
2. Forewing pattern with or without median fas-
cia, with a single midcell dot and two dots near
distal end of cell (Figs. 2-7); hindwing cubitals
of equal length (Fig. 11); first flagellomere di-
lated forming a notch; apex of uncus rounded
gnathos with two medial dentitions, proximal
flange without dilation (Fig. 12); aedeagus
short. slishtly-anecled (Fig: 13)) 22 42-2...
Pleas Aiewehahe, a e.2 eacacipsesqaine! SIs: B. normalis Meyrick
— Forewing pattern without median fascia and
with a single midcell dash and one dot on distal
end of cell near cubitus (Fig. 1); hindwing cub-
itals of unequal length (Fig. 10); first flagello-
mere not dilated; apex of uncus obtuse, gnathos
without dentitions, proximal flange with an an-
gular dilation (Fig. 8); aedeagus long, sickle-
Shapeda(t 29) ieeameuseuss enon C. darwini, n. sp.
3. Forewing pattern with or without median fas-
cia, with a single midcell dot and two dots near
distal end of cell (Figs. 2—7); hindwing cubitals
of equal length (Fig. 11); ostium near seventh
sternum, anterior margin of eighth sternum en-
tire, ductus seminalis near ostium, ductus bur-
sae long, signum hornlike (Fig. 15)
ath Re CeO Ra Ronen Pet oe B. normalis Meyrick
— Forewing pattern without median fascia, with
a single midcell dash and one dot on distal end
of cell near cubitus (Fig. 1); hindwing cubitals
of unequal length (Fig. 10); ostium near eighth
sternum, anterior margin of eighth sternum
notched, ductus seminalis not close to ostium,
349
ductus bursae short, signum platelike (Fig. 14)
3.01 8:0 °a0),5, 88 SERRA C. darwini, n. sp.
Calosima darwini Adamski and Landry,
new species
(Figs. 1, 8-10, 14)
Diagnosis.—Forewing with a midcell
dash; male genitalia with apex of uncus ob-
tuse, ventrally keeled; dorsal part of proxi-
mal flange dilated; juxta divided; aedeagus
long and sickle-shaped; female genitalia
with widened base of ductus seminalis, and
signum keeled longitudinally.
Head: Vertex and frontoclypeus with
white scales intermixed with tricolored
scales, basal %4 white, pale grayish brown
distally, with a white apex; some specimens
have mostly tricolored scales on vertex in-
termixed with white scales; outer surface of
labial palpus pale grayish brown intermixed
with white scales, mostly white to near dis-
tal margin of segments, inner surface white
intermixed with few pale grayish-brown
scales; antennal scape and pedicel white,
flagellum pale brown; proboscis white in-
termixed with pale grayish-brown scales.
Thorax: Tegula and mesoscutum white
intermixed with tricolored scales, or mostly
pale grayish-brown scales intermixed with
few white scales. Legs with outer surface
brown or pale-brown scales intermixed with
white scales. Scales white to near distal
margin of segments; inner surface of legs
white. Forewing (Fig. 1): length 4.0—6.9
mm (n = 24), mostly white intermixed with
tricolored scales, or both types of scales in-
termixed about equally; basal % of radius
and cubitus demarcated by brown scales, a
brown dash about midcell, and a brown dot
within distal part of cell near cubitus, (these
markings may be faint or absent in some
rubbed specimens); submarginal scales
mostly pale brown or brown, marginal
scales mostly white; undersurface uniform
pale brown. Venation (Fig. 10), with M,
arched and cubitals originating from com-
mon area. Hindwing with both surfaces uni-
form pale brownish gray; venation (Fig. 10)
350
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Pigs ie
Holotype of Calosima darwini.
with CuA, branched from M, near base, M,
broadly arched toward M,.
Abdomen: White.
Male Genitalia (Figs. 8-9): Uncus ven-
trally keeled, apex obtuse, slightly nar-
rowed basally, with two pairs of ventral se-
tae; base of uncus with two small depres-
sions; dorsal margin of gnathos slightly
rounded; distal %4 of digitate upper division
of valva setose, lower division basally wid-
ened, acuminate; dorsal part of proximal
flange dilated, dilation angular and setose;
juxta divided forming two semicircular
bands; vinculum narrow, medially acumi-
nate; aedeagus long and slender, sickle-
shaped, slightly bulbous basally; anellus
with microsetae.
Female Genitalia (Fig. 14): Ovipositor
telescopic, in three membranous subdivi-
sions; ostium near anteriorly notched eighth
sternum; antrum membranous and undiffer-
entiated from ductus bursae; ductus semin-
alis widened basally, narrowed distally; cor-
pus bursae with large platelike signum, lon-
gitudinally keeled and narrowed posterior-
ly.
Holotype.—d, ““ECUADOR, GALA-
PAGOS, San Cristobal, 2 km, SW P[uer]to
Baquarizo [sic], 11/II/1989, M[ercury]
V[apor] Lfight], B[ernard] Landry”. De-
posited in the Canadian National Collection
[CNC]. Specimen not dissected.
Paratypes.—Floreana: 3 d, ““ECU[ADOR],
GALAPAGOS, Floreana Punta Cormoran,
21/1V/1992, MVL, leg. B. Landry’’, speci-
men not dissected; 2 2 same data as above
except; “‘23/IV/1992’’, specimens not dis-
sected. Rabida: | ¢d, 1 2, ““ECU[ADOR],
GALAPAGOS, Rabida, Tourist trail MVL,
3/IV/1992, leg. B. Landry”’, specimens not
dissected. San Cristobal: 2 2, “ECUA-
DOR, GALAPAGOS, San Cristobal, 2km
SW P[uer] to Baquarizo [sic], 11/11/1989,
MVL, B. Landry’’, specimens not dissect-
ed; 1 2 with same data as above except;
17/11/1989”, “‘2 genitalia slide by D.
Adamski 3312” [green label], ““2 wing
slide by D. Adamski 3350” [green label].
VOLUME 99, NUMBER 2
Figs. 2-7. Adults of Blastobasis normalis, variation.
Santa Cruz: 3 6; “ECUADOR, GALA-
PAGOS, Santa Cruz, Arid Zone, 17/1/1989,
MVL, B. Landry’’, “d genitalia slide by D.
Adamski 3285” [green label], “‘d genitalia
slide by D. Adamski 3293” [green label];
1 3d paratype not dissected; 6 2; 1 & with
same data as above, [specimen not dissect-
ed]; 2 2 with same data as above except:
352
*16/1/1989"", “2 genitalia slide by D.
Adamski 3287” [green label]; “*29/1/1989”
[specimen not dissected]; 3 &:
‘““ECU[ADOR], GALAPAGOS, Santa
Cruz, ECC. De 4/1/1992, MVL, Jes: BE:
Landry” ‘“‘? genitalia slide by D. Adamski
3286” [green label]; 1 2 with same data as
above except; “‘6/III/1992, UVL’, specimen
not dissected; 1 2 with same data as above
except; “Conway, 14/IV/1992’’, “2 geni-
talia slide by D. Adamski 3288” [green la-
bel]. 5 2 specimens collected by R. Silber-
glied with the following data: ““GALAPA-
GOS I[SLANDS], SANTA CRUZ; Acade-
my Bay, C[harles] Darwin Res[earch]
Sta[tion], Aug[ust] [19]70, Altitude] (+,
=) om, JRe Silbenslied’ > ae 157 UNV
blacklight’’, “‘2 genitalia slide by D.
Adamski 615” [green label]; 3 2 with same
data as above except; ‘2 genitalia slide by
D. Adamski 616” [green label], “2 wing
slide by D. Adamski 843” [green label];
‘© genitalia slide by D. Adamski 617”
[green label], ““2 wing slide by D. Adamski
717” [green label]; one 2 specimen with
same data as above except; “25 FEB[ruary]
[19]70”, ““@ genitalia slide by D. Adamski
Gls’ encens abel santiaeo. sl) ice
““ECU[ADOR], GALAPAGOS, Santiago,
Bahia Espumilla, 4/TV/1992, MVL, leg. B.
Landry’’, ““@ genitalia slide by D. Adamski
3318” [green label]. 1 6 and 1 2 paratype
are deposited in the National Museum of
Natural History, Smithsonian Institution,
[USNM] and The Natural History Museum,
London [BMNH]. The five paratypes col-
lected by R. Silberglied are deposited in the
Museum of Comparative Zoology, [MCZ]
Harvard University. The remaining para-
types are deposited in the Canadian Nation-
al Collection [CNC], Ottawa and the per-
sonal collection of B. Landry.
Remarks.—This is the first record of Ca-
losima beyond North America. Calosima
darwini and C. lepidophaga (Clarke) are
the only known Holcocerini that share a di-
vided juxta.
Etymology.—Calosima darwini is
named after Charles Darwin, whose obser-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
vations of the flora and fauna of the Gala-
pagos Islands are fundamental to evolution-
ary thought.
Blastobasis normalis Meyrick, 1918,
new combination
(Figs. 2-7, 11-13, 15)
Auximobasis normalis Meyrick, 1918:2:
160.—Clarke, 1963:4:481.—Heppner,
1984:41.
Blastobasis crotospila Meyrick, 1926:74:
278.—Linsley and Usinger, 1966:33(7):
164.—Parkin et al., 1972:48(2):105.—
Heppner, 1984:41. New synonymy.
Diagnosis.—Male genitalia with lower
division of valva with long marginal setae;
female genitalia with moderately dense mi-
crotrichia on membrane near ostium.
Head: Frontoclypeus and vertex variable;
from grayish-brown scales tipped with
white, to nearly white; darker specimens
with scales in area between dorsal and ven-
tral margins of antennal sockets with great-
er area of white or pale grayish brown on
distal end, forming a transverse color band
across frontoclypeus; outer surface of labial
palpus brown or grayish brown intermixed
with pale grayish-brown scales tipped with
white, and few white and brown scales,
each segment white to near distal margin;
inner surface white or mostly white inter-
mixed with pale-brown scales; antennal
scape and pedicel patterned as above, male
first flagellomere dilated, forming a notch,
flagellum grayish brown; proboscis pale
grayish brown intermixed with white.
Thorax: Tegula, brown or grayish brown,
grayish brown basally, pale grayish brown,
or white distally; mesoscutum patterned as
tegula except, scales tipped with pale gray-
ish brown or white form a broad transverse
color band. Legs: outer surface with scales
grayish brown or brown tipped with white,
white near midtibia and distal margin of tar-
someres; inner surface mostly white inter-
mixed with pale grayish-brown scales.
Forewing (Figs. 2—7): length 4.2-6.4 mm
(n = 129), highly variable; grayish brown,
VOLUME 99, NUMBER 2
10
Figs. 8-10. Calosima darwini. 8, Genital capsule. 9, Aedeagus. 10, Wing venation. Figs. 8, 9, Line Scale
= 0.5 mm; Fig. 10, Line scale = 1.0 mm.
pale grayish brown or brown, most scales
tipped with white; median fascia present or
absent, complete or incomplete; one dot
near midcell and two dots near distal mar-
gin of cell; some specimens with a brown
basal streak on posterior margin (Figs. 4,
6—7); marginal dots demarcating radial, me-
dial, and cubital veins; female specimens
usually darker than males; undersurface
brown, pale brown basally; venation (Fig.
11) with M, and CuA, not strongly arched.
Hindwing with both surfaces pale grayish
354 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 11-13. Blastobasis normalis. 11, Wing venation. 12, Genital capsule. 13, Male aedeagus. Fig. 11, Line
scale = 1.0 mm; Figs. 12, 13; Line Scale = 0.5 mm.
brown; venation (Fig. 11) with cubitus
4-branched with M2, M3 stalked near mid-
length of preceding vein.
Abdomen: White.
Male genitalia (Figs. 12—13): Uncus par-
allel-sided with a rounded apex, sparsely
setose; gnathos with a pair of small denti-
tions; tergal setae present; digitate upper di-
vision of valva slightly narrowed medially,
broad at base, ventral margin setose; outer
VOLUME 99, NUMBER 2
margin of proximal flange setose; diaphrag-
ma densely microtrichiate to margins of
proximal flange; lower division of valva
with long marginal setae; juxta bandlike;
vinculum wide; aedeagus slightly angled,
with several moderately stout anellar setae.
Female genitalia (Fig. 15): Ovipositor
with four membranous subdivisions; ostium
in membrane near posterior margin of sev-
enth sternum; ostial membrane microtri-
chiate; two shallow and dentate invagina-
tions within membrane near posterolateral
margin of seventh sternum; antrum mem-
branous, short, forming a common incep-
tion for ductus seminalis and ductus bursae;
ductus bursae long with two rows of plate-
like sclerotizations within anterior end; cor-
pus bursae membranous, signum hornlike.
Types.—Auximobasis normalis Meyrick,
Lectotype 6, designated by Clarke (1963).
‘“‘Lectotype”’ [disc label], “‘Huigra, 4500
f[ee]t, Ecuador, Parish, 6—14, Lectotype,
Auximobasis normalis Meyrick, JFGC [lar-
ke], 1948” [hand-written label], ““d geni-
talia on slide 5-X-1948, JFGC[larke] 8077”
[hand-written label], ““Auximobasis normal-
is Meyr., 15/1, E. Meyrick det. in Meyrick
Collfection]’’ [hand-written label], “‘nor-
malis Meyr.” [hand-written label], “‘Meyr-
ick Coll[ection], BM 1938-290”. Blasto-
basis crotospila Meyrick, Holotype ¢d,
“Type, HT” [disc label], “‘James Island,
Galapagos, at light, sea level, 26-7-[19]24,
S[ain]t George Exped[itio]n, CL Collenet-
te’, “‘Brit{ish] Mus[eum] 1925-488”’,
““M-48”, ““Blastobasis crotospila Meyr., Tr.
Ent. Soc, Londsep. 2781926), Type!.d
[hand-written label]. ““‘BM <¢ genitalia slide
no. BM 27204’. Specimen is missing api-
cal portion of both antennae, right labial
palpus, right foreleg, and both midlegs.
Other specimens Examined.—Espanola:
1 dg, “ECUADOR, GALAPAGOS, Espan-
ola, Bahia Manzanillo, 25/1V/1992, MVL,
leg. B. Landry’’, “‘d genitalia slide by D.
Adamski 3304’ [green label], 1 @ with
same data as above except, “‘29/IV/1992”’,
““® genitalia slide by D. Adamski 3303”
[green label]. Fernandina: 1 6, 4 2,
355
“ECU[ADOR], GALAPAGOS, Fernandi-
na, Punta Espinosa, 12/V/1992, MVL, leg.
B. Landry’’; 1 2 with same data as above
except, ““2 genitalia slide by D. Adamski
33081.) jereen, label]. Floreana: 1.62
‘“FECU[ADOR], GALAPAGOS, Punta Cor-
moran, 21/I1V/1992, MVL, leg. B. Landry’’;
1 36, 2 2 with same data as above except,
~23/IV 1992 . Genovesa: 2d... “ECU
[ADOR], GALAPAGOS, Genovesa, Bahia
Darwin, 26/III/1992, MVL, leg. B. Lan-
dry” “‘d genitalia slides by D. Adamski
330577 and, 330672 ereeny labels]; 2 2:
same data as above except, *25/III/1992”’;
7 2, same data as above except, “@ geni-
talia slide by D. Adamski 3301”’, “‘3302”’,
and ‘*3307”’ [green labels]. Isabela: 2 6, 1
2, “ECUADOR, GALAPAGOS, Isla Isa-
bela, Puerto Villamil, 2/III/1989, MVL, B.
Landry’, ““d genitalia slide by D. Adam-
ski, 31574 ieteenmiabell]:, 1 ci. % likin: IW
Puerto Villamil, 3/11/1989”; ¢, 2 2, “2
km W Puerto Villamil, 5/III/1989”, ““@
genitalia slide by D. Adamski 3313” [green
label]; 2 @, “3 km N S[an]to Tomas
Agr[ricultural] Zone, 8/III/1989, MVL, B.
Landry’’, “‘2 genitalia slide by D. Adamski
3319” \[sreenlabell]ie3.eoe5 i) kim aN
P[uer]to Villamil, 9/11/1989”; 1 3, 1 &,
8.5 km N P[uer]to Villamil, 11/III/1989”’,
and... 13/TM/19897" 2) Ot lasuss Cove
13/V/1992”; 3 2, “V[olcan] Darwin, 300m
elev[ation], 15/V/1992”; 2 2, “V[olcan]
Darwin, 1240m elev{[ation], 19/V/1992”’,
‘© genitalia slide by D. Adamski 3315”
[green label]. 1 2, ““V[olcan] Darwin 630m
elev[ation], 17/V/1992”’. Marchena: 2 6,
‘““ECU[ADOR], GALAPAGOS, Marchena,
MVL,.. 12/1/1992. lee? -B. Landry’; 1-2
with same data as above except, “12/III/
1992’’, *“*@ genitalia slide by D. Adamski
33.1167°s iereen label. ePintas... thoy;
‘“ECU[ADOR], GALAPAGOS, Pinta,
13/11/1992, Plaja Ibbeston, MVL, leg. B.
Landry’; 1 6, 1 ¢, same data as above
except, 4 M1992 2.6. ISM 1992,
arid zone’’, “dé genitalia slide by D. Adam-
ski, 330977 [iercen label; 1. 9, 16/I/ 1992.
Z00mucleviationi a: 29. LA/IILL 992.
356
Figs. 14-15.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Li REPEAL ES TH
CUavveagsrssats
Female genitalia. 14, Calosima darwini. 15, Blastobasis normalis. Line scale = 1.0 mm.
VOLUME 99, NUMBER 2
400m elev{[ation]’’, “Wing slide by D.
Adamski, 3351” [green label]; 1 ¢, “18/
III/1992, 400m elev{ation]”’, ““d genitalia
slide by D. Adamski 3310” [green label];
2 6; “21/1/1992, (+, —) 15m elev{fation]’’.
San Cristébal: 2 2, ““ECUADOR, GALA-
PAGOS, San Cristobal, 2 km SW P[uer]to
Baquarizo [sic], 11/1/1989, MVL, B. Lan-
dry”. The following specimens with same
data as above except, 5 2, “14/II/1989”’,
PATMWVN9S9"s 2 O51 8/M/ 1989". 22/1i/
1989”; 3 3, “4 km SE P[uer]to Baquarizo
[sic], 12/11/1989”, **3 genitalia slide by D.
Adamski 3311” [green label]; 4 d, ““1 km
SE El Progreso, 14/II/1989°’, ‘3 genitalia
slide by D. Adamski 3300” [green label];
ore ipampaszone.. 15/11/1989" 3 1 id),
‘‘P[uer]to Baquarizo [sic], 17/II/1989”; 2
36, ‘“‘pampa zone, 18/II/1989’’; 3 3, “4 km
SE P[uer]to Baquarizo [sic], 20/11/1989”; 2
3, “‘base of Cerro Pelado, 22/11/1989”’, “d
genitalia slide by D. Adamski 3356” [green
label]; 1 ¢, “2 km SW P{[uer]to Baquarizo
[sic], 11/11/1989”, “3d genitalia slide by D.
Adamski 3299” [green label]; 1 d, same
data as above except, “20/II/1989”’. Santa
Criz: 6 6, 2 2, “ECUADOR, GALAPA-
GOS, 4 km N Puerto Ayora, 20/1/1989,
MVL, B. Landry”’ “6 genitalia slide by D.
Adamski 3292” and ‘*3158”’ [green labels],
‘*“® genitalia slide by D. Adamski 3289”’,
and ‘‘3290’’ [green labels]; 1 6, “Los Ge-
melos, 31/1/1989”; 2°6, 9 2, “Tortuga
Res[erve] W S[an]ta Rosa, 6/II/1989,
MVL, B. Landry’’, “‘d genitalia slides by
D. Adamski 3296”, and “3298” [green la-
bels], “2 genitalia slides by D. Adamski
SiS9 > 3291 “and 32977 fereen labels];
SRS? oo 2 km W' Bella “Vista, 27/11/
1989”, “3 genitalia slide by D. Adamski
3294” [green label], “2 genitalia slide by
D. Adamski 3295” [green label]; 1 d, 1 °,
“Finca S Devine, 17/III/1989”’. Santiago: 3
3, “ECU[ADOR], GALAPAGOS, Santia-
go, Bahia Espumilla, 4/1V/1992, MVL, leg.
B. Landry’; 2 6, 4 2, same data as above
exeept; 200m ‘eleviation], 5/1V/1992""*; 2
3,1 @, ““Aguacate, 520m elev[ation], 6/IV/
WOOF err WIV /1992""; and)“ 1/TV/1992
357
3, “Central, 700m elev[ation], 9/IV/1992”’;
(egs2) 2. Gerro Inn, 28/1/19927%
Remarks.—Blastobasis normalis is not
endemic to the Galapagos Islands as once
thought. This species is now known to oc-
cur on the Ecuadorian mainland as well.
DISCUSSION
Adamski (in Adamski and Brown, 1989)
provided a phylogenetic classification for
the North American Blastobasinae that in-
cluded evidence for the monophyly of both
Blastobasis Zeller and Calosima Dietz.
Blastobasis normalis shares all synapomor-
phies typical for the genus except the pos-
terior lobe of the corpus bursae. Similarly,
Calosima darwini shares all synapomor-
phies for the genus except an emarginate
juxta, valva with secondary articulatory
process, and absence of the proximal
flange. Whether Calosima darwini is en-
demic to the Galapagos is questionable.
This question can be reasonably addressed
only after the fauna of Central and South
America is better known.
ACKNOWLEDGMENTS
We thank K. Sattler, M. Shaffer and K.
Tuck of The Natural History Museum, Lon-
don, England, for loan of the lectotype of
Auximobasis normalis Meyrick and _ holo-
type of Blastobasis crotospila Meyrick. The
late Robert Silberglied provided specimens
of Calosima darwini that he collected from
the Galapagos Islands. We also thank the
authorities of the Ecuadorian Ministry of
Agriculture, the Galapagos National Park,
and the Charles Darwin Research Station
for kindly allowing field work and provid-
ing logistical support to B. Landry in 1989
and 1992. Joyce Cook, Moraima Inca, Ri-
cardo Palma, Stewart B. Peck, Bradley J.
Sinclair and Edwardo Vilema were excel-
lent and most helpful companions. Finally,
we are extremely grateful to Stewart B.
Peck, Carleton University, Ottawa, Canada,
for providing financial support for field
work and costs of reprints through an op-
erating grant for field research on arthropod
358
evolution from the Natural Sciences and
Engineering Research Council of Canada.
LITERATURE CITED
Adamski, D., and R. L. Brown. 1989. Morphology
and systematics of North American Blastobasidae
(Lepidoptera: Gelechioidea). Mississippi Agricul-
tural Forest Experiment Station Technical Bulletin
165 (Mississippi Entomological Museum No. 1),
70 pp.
Beebe, W. 1923. Notes on Galapagos Lepidoptera.
Zoologica (New York) 5: 50—59.
Butler, A. G. 1877. Lepidoptera. /n Giinther, A., ed.,
Account of the zoological collection made during
the visit of H.M.S. “Peterel’’ to the Galapagos
Islands. Proceedings of the Zoological Society of
London, 1877: 86-91.
Clarke, J. EF G. 1941. The preparation of slides of the
genitalia of Lepidoptera. Bulletin of the Brooklyn
Entomological Society 36: 149-161.
. 1955-70. Catalogue of the Type-Specimens
of Microlepidoptera in the British Museum (Nat-
ural History) described by E. Meyrick. 8 Vols.
[Vol. 4. 1963. 521 pp.]. British Museum (Natural
History, London).
Hayes, A. H. 1975. The larger moths of the Galapagos
Islands (Geometroidea, Sphingoidea & Noctuo-
idea). Proceedings of the California Academy of
Sciences (San Francisco) 40: 145-208.
Heppner, J. B. 1984. Jn Heppner, J. B., ed., Atlas of
Neotropical Lepidoptera. Checklist: Part 1. Mi-
cropterigoidea-Immoidea. Dr. W. Junk Publishers,
The Hague, The Netherlands, xiv + 112 pp.
Heppner, J. B. and B. Landry. 1994a. A new Tebenna
species from the Galapagos Islands (Lepidoptera:
Choreutidae). Tropical Lepidoptera 5(2): 123-
1253
. 1994b. A new sun moth from the Galapagos
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Islands (Lepidoptera: Heliodinidae). Tropical Lep-
idoptera 5(2): 126-128.
Hodges, R. W. In press. Gelechioidea. Jn Kristensen,
N. P, ed., Handbuch der Zoologie.
Kornerup, A. and J. H. Wanscher. 1978. Methuen
Handbook of Colour. 2nd ed. Methuen and Co..,
Ltd., London, 243 pp.
Landry, B. 1993. Additions to the knowledge of the
Pterophoridae (Lepidoptera) of the Galapagos ar-
chipelago, Ecuador, with descriptions of two new
species. Zoologische Mededelingen (Leiden) 67:
473-485.
Landry, B. and C. Gielis. 1992. A synopsis of the
Pterophoridae (Lepidoptera) of the Galapagos Is-
lands, Ecuador. Zoologische Verhandelingen (Lei-
den) 276: 1—42.
Landry, J.-E and B. Landry. 1994. A technique for
setting and mounting microlepidoptera. Journal of
the Lepidopterists’ Society 48: 205—227.
Linsley, E. G. 1977. Insects of the Galapagos (Sup-
plement). Proceedings of the California Academy
of Sciences (San Francisco), 33: 113-196.
Linsley, E. G. and R. L. Usinger. 1966. Insects of the
Galapagos Islands. Proceedings of the California
Academy of Sciences, 4th series 33(7): 113-196.
Meyrick, E. 1894. On a collection of Lepidoptera
from upper Burma. Transactions of the Entomo-
logical Society of London 1894: 1-29.
1918. Exotic Microlepidoptera. Vol. 2, 640
PP.
. 1926. Microlepidoptera from the Galapagos
Islands and Rapa. Transactions of the London En-
tomological Society 74: 269-278.
Parkin P., D. T. Parkin, A. W. Ewing, and H. A. Ford.
1972. A report on the arthropods collected by the
Edinburgh University Galapagos Islands Expedi-
tion, 1968. The Pan-Pacific Entomologist 48(2):
100-107.
Schaus, W. 1923. Galapagos Heterocera with descrip-
tions of new species. Zoologica (New York) 5:
23-48.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 359-373
STUDIES IN AQUATIC INSECTS XII: DESCRIPTIONS OF NINETEEN NEW
SPECIES OF THE GENUS OCHROTRICHIA MOSELY (TRICHOPTERA:
HYDROPTILIDAE) FROM MEXICO AND CENTRAL AMERICA
JOAQUIN BUENO-SORIA AND SILVIA SANTIAGO-FRAGOSO
Departamento de Zoologia, Institutio de Biologia UNAM, Apartado. Postal No. 70-153,
México, 04510 D.E, México.
Abstract.—Nineteen new species of the genus Ochrotrichia Mosely from Mexico and
Central America are described and the male genitalia figured: O. ixcateopana (Mexico);
O. crucecita (Panama); O. arriba (Panama); O. zihuaquia (Mexico); O. blanca (Belize);
O. maycoba (Mexico); O. eyipantla (Mexico); O. glabra (Panama); O. hondurenia
(Belize); O. corneolus (Panama); O. anomala (Panama); O. bicaudata (Panama); O.
regina (Panama); O. lupita (Panama); O. maya (Mexico); O. palmata (Mexico); O.
serrana (Mexico); O. poblana (Mexico); and O. velascoi (Mexico). This genus has a
wide distribution, from southern Canada south throughout Mexico and Central America,
the West Indies, and South America.
Key Words: taxonomy, Ochrotrichia, Trichoptera, Hydroptilidae, Mexico, Central Amer-
1ca
The genus Ochrotrichia Mosely of the
Neotropical Region has been studied by
Mosely (1937), Denning and Blickle
(1972), Bueno and Santiago (1981, 1992)
and Flint (1972). However, in Mexico, Cen-
tral, and South America there are still many
undescribed species.
This genus was previously divided into
the subgenera Ochrotrichia and Metrichia
by Flint (1972) because of his belief that
the larvae of both genera were indistin-
guishable, but the adults were quite distinct.
However, Wiggins (1996) found a number
of characters which separate larvae of Och-
rotrichia from Metrichia and considered
them as distinct genera; we follow this con-
cept. The adult of Ochrotrichia lacks an
apical spur on the foreleg, and the male
lacks modifications on the abdomen and
possesses genitalia with a large, often ex-
tremely modified tenth tergum and a rather
simple phallus. Metrichia has an apical spur
on the foreleg, the abdomen often possesses
internal sacs and/or setal brushes, and the
male genitalia has the tenth tergum reduced
to a simple membranous lobe, a button-like
cercus, dorsolateral hooks, and a very large
phallus, usually with two large hooks.
Most of the species of Ochrotrichia here
described, were provided by Dr. Oliver S.
Flint, Jr. from the collection of the National
Museum of Natural History (NMNH)
where they will be deposited. The rest of
the species, will be deposited in the collec-
tion of the Instituto de Biologia UNAM
(IBUNAM).
Family Hydroptilidae
In Mexico, Central, and South America
there are representatives of all six tribes:
Leucotrichiini, Stactobiini, Neotrichiini,
Ochrotrichiini, Orthotrichiini and Hydrop-
tiliini all in the subfamily Hydroptilinae
(Marshall 1979). The genus Ochrotrichia,
360
belonging to the tribe Ochrotrichiini, is well
represented in the tropical rain forests of
Mexico, Central, and South America
(Bueno and Santiago 1992).
Flint (1972), divided the genus Ochrotri-
chia Mosely into two major groups, one,
the xena group, which is characterized by a
simple ninth segment and the tenth tergum
being a simple flap often with a few small
spines. The other major group is character-
ized by the ninth tergum depressed below
the dorsal margin of the pleura and the an-
terior margin is produced beyond the ante-
rior margin of the pleura. The tenth tergum
is more or less fused with the ninth tergum
and appears greatly modified, bearing many
spines, processes and plates. This second
major group was divided by Flint (1972)
into six subgroups, on the basis of the shape
of the tenth tergum, inferior appendages
and phallus.
Genus Ochrotrichia Mosely
Polytrichia Sibley 1926:102.—Betten
1934:153.—Mosely 1937:184.—Ross
1944:125. Type species: Ithytrichia con-
fusa Morton 1905, monobasic [preocu-
pied].
Ochrotrichia Mosely 1934:162.—Ross
1944:125.—Denning 1956:255.—Flint
1964:58; 1968a:48; 1968b:59. Type spe-
cies Ochrotrichia insularis Mosely, 1934,
by original designation.
Ochrotrichia ixcateopana Bueno and
Santiago, new species
(Figs. 1-3)
On the basis of the shape of the inferior
appendages in lateral view and because of
the presence and distribution of the pro-
cesses of the tenth tergum, O. ixcateopana
belongs to the group of species related to
O. dactylophora Flint, from which it differs
in the long process of the inferior append-
ages and features of the tenth tergum.
Adult.—Length of forewing 2.5 mm.
Color in alcohol dark brown. Male genita-
lia: ninth tergum depressed and produced
anteriad. Tenth tergum with two spines on
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
right side, lateralmost shorter, and other
long and curved ventrally; centrally with
basal middorsal process having two up-
turned spines at apex. Inferior appendages
asymmetrical, both with heavy, black, pos-
terior spines; left inferior appendage with
row of three short spines in midsection of
posterior border; right inferior appendage
with two, black, short, robust spines. Phal-
lus long and threadlike.
Material.—Holotype, ¢6: MEXICO:
Guerrero, 7 km Route Taxco-Ixcateopan, 17
June 1987, J. Bueno, H. Brailovsky and E.
Barrera (IBUNAM).
Etymology.—Named for Ixcateopan,
Guerrero, the area where the type was col-
lected.
Ochrotrichia crucecita Bueno and
Santiago, new species
(Figs. 4—5)
This species appears to be closely related
to Ochrotrichia cruces Flint. However, it is
distinguished from that species by the pres-
ence of two spines on the right side of the
tenth tergum in lateral view: one being
short, stout, basodorsal, curved ventrally
and the other long and curved ventrally, and
by the lack of the third small, thin, spine
on the right side, which is present in O.
cruces.
Adult.—Length of forewing 3 mm. Color
in alcohol dark brown. Male genitalia: ninth
segment with short, broad posterodorsal
lobe, in lateral aspect deeply depressed and
produced anteriad. Tenth tergum complex
in dorsal view, with an elongate, black-
tipped process arising basolaterally on right
side curving to left side; a short, black-
tipped, middorsal, upcurved spine and a ro-
bust spine situated dorsolaterally; apical
process with tip hooked ventrad; in lateral
aspect elongate, black-tipped process ap-
pears curved and ventrally situated to rest
of structures; in this view, a short, black-
tipped, middorsal, spine appears situated
behind a robust, dorsolaterally situated
spine. Inferior appendages broadly trian-
gular, apex rounded, with a band of black,
VOLUME 99, NUMBER 2
Figs. 1-5.
peglike setae apically and on midbasal
ridge, one more cluster near dorsal margin.
Phallus long and threadlike.
Material.—Holotype, 6: PANAMA:
Chiriqui, Guadalupe Arriba 8°52'26’N:
82°33'13"W, 2-8 May 1984, H. Wolda
(NMNH). Paratypes: same as holotype but
3-9 July 1985, | 6 (NMNH); same as ho-
lotype but 24—30 July 1985, 1 d (NMNH).
Etymology.—crucecita: diminutive in
Spanish for little cross.
Ochrotrichia arriba Bueno and Santiago,
new species
(Figs. 6—7)
On the basis of the shape of the inferior
appendages in lateral aspect, this species
could be considered related to Ochrotrichia
1-3, Ochrotrichia ixcateopana, male genitalia. 1, Right side. 2, Left side. 3, Dorsal. 4—5, O.
crucecita, male genitalia. 4, Dorsal. 5, Lateral.
aldama (Mosely). However, the tenth ter-
gum of O. arriba is formed by two short,
rounded plates in dorsal view, a character
distinctly different from O. aldama.
Adult.—Length of forewing 3 mm. Col-
or in alcohol dark brown. Male genitalia:
ninth tergum deeply depressed and pro-
duced anteriad. Tenth tergum in dorsal as-
pect like a simple, broad, elongate flap
with a rounded, ringlike area anteriad and
a curved and narrowed posterior plate; in
lateral aspect apex of posterior plate ap-
pears curved ventrad. Inferior appendages
in lateral view rectangular, broad and with
small posteroventral lobe; posterior margin
with a band of black, peglike setae. Phallus
long and threadlike.
Material.—Holotype, ¢: PANAMA:
362
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 6-10. 6—7, Ochrotrichia arriba, male genitalia. 6, Dorsal. 7, Lateral. 8-10, O. zihuaquia, male genitalia.
8, Right side. 9, Left side. 10, Dorsal.
Chiriqui, Guadalupe Arriba, 8°52'26"N
82°33'13"W, 3-9 July 1985, H. Wolda
(NMNH).
Etymology.—arriba: Spanish for above.
Ochrotrichia zihuaquia Bueno and
Santiago, new species
(Figs. 8-10)
On the basis of the shape of the inferior
appendages, this species appears to be related
to Ochrotrichia insularis Mosely, but the dis-
tinct processes on the tenth tergum, resemble
those of Ochrotrichia aldama (Mosely).
Adult.—Length of forewing 2.9 mm.
Color in alcohol dark brown. Forewing
with a transverse, white band at midlength.
Male genitalia: ninth segment open dorsal-
ly, slightly produced anteriad. Tenth tergum
mostly membranous with three long pro-
cesses almost of equal length; in lateral as-
pect apparently with two long processes
with acute apices; process on left side with
a group of small spines. Inferior append-
ages short and wide with cluster of thick,
long, dark spines on mesal face at apex; in
dorsal aspect, appendages asymmetric, left
appendage with long, apicodorsal process,
right appendage with posterior margin
swollen and covered with peglike seta.
Phallus long and threadlike.
Material.—Holotype, d6: MEXICO: Guer-
rero, route 134, 102 km N. W. of Zihuatanejo,
1200 m, 7 June 1984, J. Bueno and E. Ma-
rino (IBUNAM). Paratypes: MEXICO: Oa-
xaca, Candelaria de Loxicha, Finca Pacifica,
510 M elevation, 2 June 1985, E. Barrera 1
36 (BUNAM) and 1 d (NMNH).
Etymology.—Named from the place
where the type was collected in Zihuaquio,
Guerrero.
VOLUME 99, NUMBER 2
Ochrotrichia blanca Bueno and
Santiago, new species
(Figs. 11—13)
This species and the two subsequent spe-
cies, on the basis of shape of the inferior
appendages in lateral aspect and the pres-
ence of long processes on the right side of
the tenth tergum, appear to be related. How-
ever, Ochrotrichia blanca differs from these
species by the presence of a short middorsal
spine and by the distribution of the five
long processes on the tenth tergum.
Adult.—Length of forewing 2 mm. Color
in alcohol pale brown. Male genitalia: ninth
segment deeply depressed and produced an-
teriad. Tenth tergum in dorsal view, with a
long dorsalmost process on left side, with a
short middorsal spine; on right side with
two, long, curved processes, left one wider
than right one which appears narrowed,
both with apex acute and convergent; in lat-
eral view ventral plate appears U-shaped on
left side. Inferior appendages elongate,
broadest apically; in lateral aspect slightly
triangular; posterior border with many
black, peglike setae. Phallus long and
threadlike.
Material.—Holotype, 6: BELIZE: Cayo
District, Rio Privassion, Blancaneaux
Lodges -9-11 July 1973, Yo, Sedman
(NMNH).
Etymology.—blanca: feminine, white in
Spanish.
Ochrotrichia maycoba Bueno and
Santiago, new species
(Figs. 14-16)
This species appears related to Ochrotri-
chia blanca and O. eyipantla. Because of
the distribution of the processes of the tenth
tergum, and the shape of inferior append-
ages, O. maycoba also shows a relationship
with the species of the confusa complex.
Adult.—Length of forewing 3 mm. Color
in alcohol pale brown. Male genitalia: ninth
tergum deeply depressed and slightly pro-
duced anteriad. Tenth tergum with five
elongate processes; two processes on left
363
side and three on right side, dorsalmost one
appears widest and slightly curved to right;
underneath these five processes a long pro-
cess ending in a sclerotized, hook-like apex.
Inferior appendages broadly triangular, with
one broad, posterodorsal process in lateral
aspect; right appendage with a posterior,
apically situated, patch of black peglike se-
tae; left appendage with apical row of black
peglike setae and a fingerlike process on
posterodorsal corner. Phallus long and
threadlike.
Material_—Holotype, ¢: MEXICO: So-
nora, Maycoba River, west of Maycoba, 21
August 1986, R. Baumann, Sergeant &
Kondratieff (NMNH).
Etymology.—Named for the Maycoba
River where the holotype was collected.
Ochrotrichia eyipantla Bueno and
Santiago, new species
(Figs. 17-19)
On the basis of the shape and distribution
of the processes of the tenth tergum, this spe-
cies appears to be related to Ochrotrichia
maycoba. However, Ochrotrichia eyipantla
can be separated from this species by the
presence of only four spinelike processes on
the tenth tergum, and by the differences in
shape and size of the inferior appendages.
Adult.—Length of forewing 2 mm. Color
in alcohol pale brown. Male genitalia: ninth
tergum slightly depressed and slightly pro-
duced anteriad. Tenth tergum in dorsal as-
pect with a long, wide, and laterally-situ-
ated process, whose apex touches the apical
hooklike portion of ventralmost process
which is widened at midlength; with four
shorter middorsal spinelike processes, later-
almost on right side shorter than lateralmost
on left side, one of middorsal spinelike pro-
cesses slightly curved; in lateral view on
right side with a straight spinelike process.
Inferior appendages broadly triangular, with
apical row of black peglike setae. Phallus
long and threadlike.
Material.—Holotype, ¢: MEXICO: Ve-
racruz, Salto de Eyipantla, Eyipantla River,
Figs. 11-16.
O. maycoba, male genitalia. 14, Right side. 15, Left side. 16, Dorsal.
30 March 1990, P. J. Spangler and S. San-
tiago-Fragoso (IBUNAM).
Etymology.—Named from Eyipantla, a
waterfall in Veracruz, where the holotype
was collected.
Ochrotrichia glabra Bueno and
Santiago, new species
(Figs. 20-21)
On the basis of the shape of the tenth
tergum, Ochrotrichia glabra appears most
similar to Ochrotrichia hondurenia. How-
ever, O. glabra can be separated from that
species by the lack of stout peglike spines
on the tenth tergum.
Adult.—Length of forewing 2 mm. Color
in alcohol dark brown. Male genitalia: ninth
segment deeply depressed and slightly pro-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
11-13, Ochrotrichia blanca, male genitalia. 11, Right side. 12, Left side. 13, Dorsal. 14-16,
duced anteriad. Tenth tergum a simple,
elongate, glabrous lobe, with apex rounded.
Inferior appendages rectangular and elon-
gate, apex broad and rounded; with cluster
of black peglike setae at apex and another
group near ventral margin; midbasal ridge
obsolescent. Phallus long and threadlike.
Material.—Holotype, ¢d: PANAMA,
Chiriqui, Guadalupe Arriba, 8°52’26"N and
82°33'13”W, 30 October—5 November 1985,
H. Wolda (NMNH).
Etymology.—glabra: Latin for hairless,
smooth, relating to the tenth tergum.
Ochrotrichia hondurenia Bueno and
Santiago, new species
(Figs. 22—23)
On the basis of the tenth tergum, this spe-
cies appears related to Ochrotrichia glabra.
VOLUME 99, NUMBER 2
Figs. 17-19.
lia. 17, Right side. 18, Left side. 19, Dorsal.
Ochrotrichia eyipantla, male genita-
However, because of the presence of three,
stout, darkened, apically-situated, peglike
processes on the tenth tergum and two long
filaments on the apical portion of the phallus,
Ochrotrichia hondurenia is easily distin-
guished.
Adult.—Length of forewing 2 mm. Color
in alcohol pale brown. Male genitalia: ninth
segment deeply depressed and produced an-
teriad. Tenth tergum a broad, elongate plate,
with three stout, darkened, apically-situat-
ed, peglike processes; with left and right
side sclerotized, but membranous centrally.
Inferior appendages elongate, apex round-
ed; with a row of black peglike setae
around apex and along ventral margin.
Phallus tubular with a conically expanded
base; apex with two processes, one straight
and another gradually decurved.
Material.—Holotype, ¢: BELIZE: Cayo
District, Mountain Pine Ridge, 27 June
1971, G. Stacell (NMNH).
Etymology.—Named for the former Brit-
ish Honduras, the country where the holo-
type was collected.
Ochrotrichia corneolus Bueno and
Santiago, new species
(Figs. 24—25)
On the basis of the elongate shape of the
inferior appendages and the presence of one
short, strong, basodorsal spine, on the tenth
tergum, this species is related to Ochrotri-
chia stylata (Ross). However, O. corneolus
can be separated from that species by the
shape of the curved, basodorsal spine on
the tenth tergum.
Adult.—Length of forewing 2 mm. Color
in alcohol pale brown. Male genitalia: ninth
tergum with rounded posterodorsal lobe in
lateral aspect, deeply depressed and slightly
produced anteriad. Tenth tergum a narrow,
elongate plate with apex upturned; dorsally
with a short, single, sinuate, basal spine;
underneath plate on left side arises a long
spinelike process with its apex a little lon-
ger than plate. Inferior appendages elon-
gate, narrow, apex rounded with a row of
black, peglike setae apically, along the ven-
tral margin and midbasal ridge. Phallus
long and threadlike.
Material.—Holotype, ¢: PANAMA:
Barro Colorado Island, Snyder-Molino
Trail, Marker 3, Trap Level III, 18-24 No-
vember 1987, H. Wolda (USNM). Para-
types: same as holotype, 1 ¢d (NMNH);
same but 3-9 June 1987, 1 ¢d (NMNH);
same but 25 November—1| December 1987,
1 36 (NMNH); same but 2—8 December
1987, 1 6 (NMNH); same but 19-25 Oc-
tober 1988, 1 6 (NMNH); same but 10-16
May 1989, 1 6 (NMNH); same but 21—27
June 1989 1 5 (NMNH); same but 13 Sep-
tember 1989, 1 d6 (NMNH); same but 22—
28 November 1989, 1 6 (NMNH); same
but 29 November—5 December 1989, | d
(NMNH); same but 6—12 December 1989,
1 6 (IBUNAM).
Etymology.—corneolus: Latin for a di-
minutive horn.
Ochrotrichia anomala Bueno and
Santiago, new species
(Figs. 26—27)
On the basis of the shape of the inferior
appendages and tenth tergum, this species
366
Figs. 20-27.
may be related to Ochrotrichia flagellata
Flint, from which it can be separated by the
presence of sclerotized bars on the tenth ter-
gum, and by the lightly rectangular shape
of the ninth segment in lateral aspect.
Adult.—Length of forewing 2 mm. Color
in alcohol pale brown. Male genitalia: ninth
segment deeply depressed and slightly pro-
duced anteriad, in lateral aspect appearing
triangular. Tenth tergum a membranous
hood, dorsally with a pair of sclerotized,
sinuous bars. Inferior appendages long,
slender, apex rounded with a mesal cluster
of black peglike setae; a high dorsal lobe,
with a row of black peglike setae at margin
of apex. Phallus long and threadlike.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
20-21, Ochrotrichia glabra, male genitalia. 20, Right side. 21, Dorsal. 22—23, O. hondurenia,
male genitalia. 22, Dorsal. 23, Lateral. 24—25, O. corneolus, male genitalia. 24, Left side. 25, Dorsal. 26-27,
O. anomala, male genitalia. 26, Dorsal. 27, Right side.
Material.—Holotype, ¢: PANAMA:
Barro Colorado Island, Snyder-Molino
Trail, Marker 3, trap level III, 18—24 No-
vember 1987, H. Wolda (NMNH). Para-
types: same as holotype but 7—13 October
1987, 1 6 (NMNH); same but 18-25 Oc-
tober 1988, 1 d (NMNH); same but 16—22
November 1988, 1 d (NMNH); same but
23-29 November 1988, 1 3d (NMNH);
same but 8-14 February 1989, 1 ¢
(NMNH); same but 11—17 October 1989, 1
6 (NMNH); same but 7—13 November
1990, 1 6 (NMNH); same but 19-25 De-
cember 1990, 1 6 (IBUNAM).
Etymology.—anomala: means abnormal
in Spanish.
VOLUME 99, NUMBER 2
Figs. 28-37.
32, O. regina, male genitalia. 31, Right side. 32, Dorsal. 33-35, O. lupita, male genitalia. 33, Right side. 34,
Left side. 35, Dorsal. 36-37, O. maya. 36, Left side. 37, Dorsal.
Ochrotrichia bicaudata Bueno and
Santiago, new species
(Figs. 28-30)
On the basis of the shape of the inferior
appendages and the tenth tergum, this spe-
cies appears similar to Ochrotrichia aldama
(Mosely). However, O. bicaudata is rec-
ognized by the presence of two, long, slen-
der processes on the tenth tergum.
Adult.—Length of forewing 2 mm. Color
in alcohol pale brown. Male genitalia: ninth
tergum deeply depressed and produced an-
teriad in lateral aspect, with rounded pos-
terodorsal lobes. Tenth tergum with two,
long, slender processes, one appears sinu-
ous and wider with apex curving to right;
on right side a slightly-curved, long pro-
cess, with apex directed ventrad. Inferior
appendages elongate, with rounded apex;
apex, ventral margin and midbasal ridge
28-30, Ochrotrichia bicaudata, male genitalia. 28, Right side. 29, Left side. 30, Dorsal. 31—
with many, short, black, peglike setae. Phal-
lus long and threadlike.
Material.—Holotype, ¢: PANAMA:
Barro Colorado Island, Snyder-Molino
Trail, Marker 3, trap level III, 2-8 Decem-
ber 1987, H. Wolda (NMNH). Paratypes:
Same as holotype, 1 ¢ (NMNH); same but
11-17 March 1987, 1 6 (NMNH); same
but 7-13 October 1987, 1 6 (NMNH);
same but 4-10 November 1987, 1 ¢o
(NMNH); same but 18-24 November 1987,
3 ¢ (NMNH); same but 16—22 November
1988, 1 6 (NMNH); same but 30 Novem-
ber—6 December 1988, 1 d (NMNH); same
but 7-13 December 1988, 2 6 (NMNH);
same but 4—10 January 1989, 1 6
(NMNH); same but 11-17 January 1989, 1
6 (NMNH); same but 1-7 February 1989,
1 d6 (NMNH); same but 6—12 December
1989, 1 6 (NMNH); same but 13-19 De-
368 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
cember 1989, | 6 (NMNH); same but 8—
14 August 1990, 1 6 (NMNH); same but
22-28 August 1990, 1 6 (NMNH); same
but 12-18 September 1990, 1 ¢6 (NMNH);
same but 17-23 October 1990, 1 ¢
(NMNH); same but 16—22 January 1991, 1
6 (IBUNAM).
Etymology.—bicaudata: Latin, for the
two processes of the tenth tergum.
Ochrotrichia regina Bueno and
Santiago, new species
(Figs. 31—32)
Because of the membranous and elongate
aspect of the tenth tergum, this species ap-
pears related to Ochrotrichia lupita. How-
ever, the presence of one, curved, spinelike
process apically on the tenth tergum easily
distinguishes Ochrotrichia regina.
Adult.—Length of forewing 2 mm. Color
in alcohol dark brown. Male genitalia: ninth
tergum deeply depressed and produced an-
teriad; in lateral aspect appearing almost
quadrangular. Tenth tergum with short, ba-
sal, middorsal plate; apical process with tip
pointed and curved to right; in lateral aspect
with apex of apical process turned ventrally
and slightly enlarged preapically. Inferior
appendages in lateral view elongate, apex
rounded; apex, ventral margin and midbasal
ridge with with a band of black peglike se-
tae. Phallus long and threadlike.
Material.—Holotype, ¢: PANAMA:
Barro Colorado Island, Snyder Molino trail,
Marker 3, Trap level-I, 30 November—6 De-
cember 1988, H. Wolda (NMNH). Para-
types: Same as holotype, but 27 May—2
June 1987, 1 ¢ (NMNH); same but 5-11
August 1987, | 6 (NMNH); same but 26
August-1 September 1987, 1 ¢ (NMNH);
same but 16-22 September 1987, 1 ¢
(NMNH); same but 30 September—6 Octo-
ber 1987, 1 6 (NMNH) 1 6 (BUNAM);
same but 21-27 October 1987, 1 6
(NMNH); same but 4—10 November 1987
1 6 (NMNH); same but 18—24 November
1987, 1 6 (NMNH); same but 25 Novem-
ber—1 December 1987, 1 ¢ (NMNH); same
but 2—8 December 1987, 1 6 (NMNH);
same but 18-24 May 1988, 1 d6 (NMNH);
same but 9-15 November 1988, 2 ¢
(NMNH); same but 23—29 November 1988,
1 6 (NMNH); same but 26 April—2 May
1989, 1 6 (NMNH); same but 24—30 May
1989, 1 ¢d (NMNH); same but 30 August—
5 September 1989, 2 d (NMNH); same but
6-12 September 1989, 3 d (NMNH); same
but 22—28 November 1989, 2 d6 (NMNH);
same but 9-15 May 1990, 1 ¢6 (NMNH);
same but 13—19 June 1990, 1 ¢6 (NMNH);
same but 20—26 June 1990, 1 d6 (NMNH);
same but 27 June—3 July 1990, 1 ¢
(NMNH); same but 18—24 July 1990, 1 ¢d
(NMNH); same but 8-14 August 1990, 1
36 (NMNH); same but 15-21 August 1990,
1 36 (NMNH); same but 22-28 August
1990, 1 d6 (NMNH); same but 29 August—
4 September 1990, 2 d (NMNH); same but
12-18 September 1990, 1 6d (NMNH);
same but 19-25 September 1990, 1 6
(NMNH); same but 26 September—2 Octo-
ber 1990, 1 d6 (NMNH); same but 17—23
October 1990, 2 d (NMNH); same but 31
October—6 November 1990, 1 d (NMNH);
same but 14-20 November 1990, 2 6
(NMNH); same but 21—27 November 1990,
2 6 (TBUNAM).
Etymology.—regina: Latin for queen.
Ochrotrichia lupita Bueno and Santiago,
new species
(Figs. 33-35)
On the basis of the hook-shaped apex of
the tenth tergum, this species is related to
Ochrotrichia regina. However, O. lupita is
easily distinguished by the lack of a broad,
middorsal area on the tenth tergum.
Adult.—Length of forewing 3 mm. Color
in alcohol dark brown. Male genitalia: ninth
segment deeply depressed and produced an-
teriad. Tenth tergum with a short, basodor-
sal plate with a long, apical process whose
apex curves ventrad; left side in lateral
view with an elongate, spinelike process
and underneath this a long, slightly sinuous,
spinelike structure ventrally directed. Infe-
rior appendages elongate, broad, apex
rounded, with a cluster of black, peglike se-
VOLUME 99, NUMBER 2
tae at apex and another group near ventral
margin. Phallus long and threadlike.
Material.—Holotype, 6: PANAMA:
Chiriqui, Guadalupe Arriba, 8°52'26’N:
82°33'13"W, 3-9 July 1985, H. Wolda
(NMNH). Paratype: same as holotype but
1-7 May 1985, H. Wolda. 1 ¢ (NMNH).
Etymology.—lupita: nickname in Span-
ish for Guadalupe.
Ochrotrichia maya Bueno and Santiago,
new species
(Figs. 36—37)
Because the presence of two asymmetri-
cal precesses on the tenth tergum, Ochro-
trichia maya is not obviously related to any
other species of this genus.
Adult.—Length of forewing, 2 mm. Col-
or stramineous in alcohol. Male genitalia:
ninth tergum depressed and produced an-
teriad. Tenth tergum divided into long and
short processes; long process in left lateral
view clearly curved with tip directed ven-
trally; in dorsal view with basal portion
shieldlike, with three, short teeth on right;
short process wider, with tip directed ven-
trad. Inferior appendages elongate, broad;
with a band of black, peglike setae at apex
and a group of peglike setae mesad. Phallus
long and threadlike.
Material.—Holotype, ¢: MEXICO: Chia-
pas, Cascada de Misolja, 20 km. S. from
Palenque, 17-18 May 1981, C. M. & O. S.
Flint Jr. (NMNH). Paratype: Route 195, 1.5
mi. N. Ixhuatén, 23 December 1983. S.
Hamilton, R. Holzenthal, M. Kovach, 1 ¢d
(NMNH).
Etymology.—maya: Indian people living
in the area where the type was collected.
Ochrotrichia palmata Bueno and
Santiago, new species
(Figs. 38-39)
Because of the presence of short, stout,
spines at the base of the tenth tergum, Och-
rotrichia palmata may be related to Och-
rotrichia poblana and O. velascoi. How-
ever, O. palmata can be recognized by the
369
Figs. 38-39.
Ochrotrichia palmata, male genitalia.
38, Left side. 39, Dorsal.
shape of the tenth tergum which appears
palmate in lateral aspect.
Adult.—Length of forewing, 2.5 mm.
Color dark brown in alcohol. Forewing
with a transverse white band at midlength.
Male genitalia: ninth segment open dorsal-
ly, produced anteriad; posterior margin sin-
uate. Tenth tergum in lateral aspect basally
narrow, with curved midsection, apical pro-
cess palmate, with a fingerlike, ventrally-
curved process, and one, short, basal spine;
in dorsal aspect, with sinuous, membranous
process, surrounded by four short spines,
centralmost black-tipped and covered by
membranous, dorsal process. Inferior ap-
pendages, narrow, with rounded apex; apex
of midbasal ridge with a line of black, peg-
like setae. Phallus long and threadlike.
Material.—Holotype, ¢: MEXICO: Es-
370
Figs. 40-41.
40, Right side. 41, Dorsal.
Ochrotrichia serrana, male genitalia.
tado de Mexico, Temascaltepec, 8 February
1979, J. Bueno and J. Padilla IBUNAM).
Etymology.—palmata: Latin for hand-
shape, shown in the lateral aspect of the
tenth tergum.
Ochrotrichia serrana Bueno and
Santiago, new species
(Figs. 40—41)
On the basis of the shape and distribution
of the processes of the tenth tergum, this
species is similar to Ochrotrichia chiapa
Denning. However, Ochrotrichia serrana
differs from that species by the possession
of a stout spine with denticuli on the dorsal
margin and by the presence of a short,
spinelike process on the left side of the base
of tenth tergum.
Adult.—Length of forewing 2.5 mm.
Color pale brown in alcohol. Male genita-
lia: ninth segment deeply depressed and
produced anteriad. Tenth tergum in dorsal
aspect with a basodorsal, stout process with
dorsal edge serrate; two lateral spines on
right side, dorsalmost wider and curved to
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
left, ventralmost long and slender, ending
with darkened apex; left side with a short,
basolateral spine with dorsal margin serrate;
long middventral process widened at mid-
length with tip rounded and curved to right.
Inferior appendages in dorsal view with
ventral margin bearing a line of peglike se-
tae; in lateral view long and slender, apex
acute. Phallus long and threadlike.
Material.—Holotype, ¢d. MEXICO:
Guerrero, Acahuizotla, 10 km E. of Chil-
pancingo, 10 November 1982, J. Bueno, E.
Barrera and H. Brailovsky (IBUNAM).
Etymology.—serrana: Spanish name, giv-
en to women inhabitants of the mountains.
Ochrotrichia poblana Bueno and
Santiago, new species
(Figs. 42—43)
On the basis of elongate inferior append-
ages, tapering to a pointed apex in lateral
view, this species is related to the /ometa
group, especially with Ochrotrichia orego-
na Ross or O. nacora Denning and Blickle.
However, O. poblana can be separated from
these species, by the presence of stout,
spinelike processes on both inferior append-
ages and, in lateral view, by the presence
on the tenth tergum of one, dorsally curved,
spinelike process on the left side, with two
apices, one directed posteriad and one di-
rected anteriad.
Adult.—Length of forewing 3 mm. Color
in alcohol dark brown. Male genitalia: ninth
segment incomplete dorsally, produced an-
teriad, with posterior border slightly sinu-
ous in lateral aspect. Tenth tergum in dorsal
aspect with a short, basodorsal spine arising
from left side, curved apically; on right side
with one, black, basodorsal spine barely
passing left spine; a stout, dark-tipped pro-
cess, attached in middle, with basal end
curved upward and directed anteriad and
distal end ventrally curved and directed
posteriad. Left inferior appendage in dorsal
aspect with a well developed midbasal,
stout, black-tipped spine; right appendage
with a short, basoventral spine and a mid-
ventral line of teeth; in lateral aspect a mid-
VOLUME 99, NUMBER 2
ventral, stout, dark-tipped spine; apex end-
ing in a short, black, stout process. Phallus
long and threadlike.
Material.—Holotype, 6: MEXICO:
Puebla, km 30, route Zacapoaxtla-Zacatlan,
May 2 1987, J. Bueno, E. Barrera (IBUN-
AM). Paratypes: same data, 1 6 (USNM)
and 1 6 (IBUNAM); Route Cuetzalan-Za-
catlan, km 64, elevation 1,250 m, March 19
1987, J. Bueno, E. Barrera and H. Brailov-
sky 26 6 (IBUNAM).
Etymology.—poblana: A name given to
the women born in Puebla, Mexico.
Ochrotrichia (O.) velascoi Bueno and
Santiago, new species
(Figs. 44—45)
Because of the wide basal portion and
narrow, elongate apical portion of the in-
ferior appendages in lateral view, this spe-
cies appears to be related to the Californian
Ochrotrichia buccata Denning and Blickle.
However, the shape and distribution of the
spinelike processes of the tenth tergum will
distinguish Ochrotrichia velascoi from O.
buccata and others.
Adult.—Length of forewing, 2.5 mm.
Color dark brown in alcohol. Male genita-
lia: ninth segment open dorsally and pro-
duced anteriad. Tenth tergum in dorsal as-
pect, with a short, middorsal, curved spine
barely projecting beyond apex of a larger,
dark-tipped spine; on right side with long,
dark-tipped spinelike process; apical pro-
cess wide, membranous, slightly sinuous
and rounded at apex, arising from apex of
shortest, middorsal, dark-tipped spine; in
lateral aspect on right longest process ap-
pears wide basally and slightly curved ven-
trad, dorsalmost, dark-tipped spine shows
only dark tip; apical process membranous,
longer than other processes, and clearly
curved ventrad. Inferior appendages in lat-
eral aspect elongate, tapering to a pointed
apex; three or more peglike seta near apex;
in dorsal aspect, mesal face of left inferior
appendage with a short, dark-tipped spine
near base; apex of both appendages with
Figs. 42—43.
lia. 42, Left side. 43, Dorsal.
Ochrotrichia poblana, male genita-
three or more peglike setae. Phallus long
and threadlike.
Material—Holotype 6: MEXICO: Guer-
rero, route 134, 102 km N. W. of Zihuata-
nejo, 1200 m, 7 June 1984, J. Bueno and
E. Marino (IBUNAM). Paratype, same as
holotype, 1 6 (USNM).
Etymology.—We dedicate this species to
our student, Biol. Hector Velasco Madrigal.
ACKNOWLEDGMENTS
We are indebted to Dr. Oliver S. Flint Jr.,
Emeritus Curator of Neuropteroids of the
Department of Entomology, National Mu-
seum of Natural History, Smithsonian In-
45
Figs. 44-45. Ochrotrichia velascoi, male genita-
lia. 44, Right side. 45, Dorsal.
stitution, for his comments on the manu-
script; to Dr. P. J. Spangler for his invitation
to work under a grant from the National
Museum of Natural History, Smithsonian
Institution; to Mrs. Phyllis Spangler for her
help with the word processing of the manu-
script; to Miss Nancy Adams, support staff
of the National Museum of Natural History,
Smithsonian Institution, for her assistance
while we were working in the Museum; to
M. en C. Enrique Marino for his help while
collecting in the field; to Biol. Rafael Barba
Alvarez, our technician, for his assistance
in preparing the specimens from the collec-
tion of the UNAM; and finally to the anon-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ymous reviewers for their time and effort
to improve the manuscript.
LITERATURE CITED
Betten, C. B. 1934. The caddis flies or Trichoptera of
New York. New York State Museum Bulletin 292:
1-576.
Bueno-Soria, J. and S. Santiago-Fragoso. 1981 (for
1980). Una Nueva Especie del Género Ochrotri-
chia Mosely (Trichoptera: Hydroptilidae) Del Es-
tado de Hidalgo, Mexico. Anales del Instituto de
Biologia Universidad Nacional Autonoma de
México, Serie Zoologia 51(1): 383-388.
1992. Studies in aquatic insects, XI: seven
new species of the genus Ochrotrichia (Ochrotri-
chia) From South America (Trichoptera: Hydrop-
tilidae). Proceedings of Entomological Society of
Washington 94: 439-446.
Denning, D. G. 1956. Trichoptera, pp. 237-270. In
Usinger, R. L., ed., Aquatic Insects of California.
University of California Press Berkeley, x + 508
Pp-
Denning, D. G. and R. L. Blickle. 1972. A review of
the genus Ochrotrichia (Trichoptera: Hydroptili-
dae). Annals of the Entomological Society of
America 65: 141-151.
Flint, O. S., Jr. 1964. The caddisflies (Trichoptera) of
Puerto Rico. Technical Paper of the Agricultural
Experiment Station, University of Puerto Rico 40:
1-80.
. 1968a. The caddisflies of Jamaica (Trichop-
tera). Bulletin of the Institute of Jamaica, Science
Series 19: 1-68.
1968b. Bredin-Archbold-Smithsonian Bio-
logical Survey of Dominica, 9. The Trichoptera
(Caddisflies) of the Lesser Antilles. Proceedings
of the United States National Museum 125(3665):
1-86.
1972. Studies of Neotropical Caddisflies,
XIII: the genus Ochrotrichia from Mexico and
Central America (Trichoptera: Hydroptilidae).
Smithsonian Contributions to Zoology 118: 1-28.
Marshall, J. E. 1979. A review of the genera of the
Hydroptilidae (Trichoptera). Bulletin of the British
Museum (Natural History), Entomology Series
39: 135-239.
Morton, K. J. 1905. North American Hydroptilidae.
Bulletin of the New York State Museum 86: 63-—
85.
Mosely, M. E. 1934. New exotic Hydroptilidae.
Transactions of the Royal Entomological Society
of London 82: 137-163.
1937. Mexican Hydroptilidae (Trichoptera).
Transactions of the Royal Entomological Society
of London 86: 151—190.
Ross, H. H. 1944. The caddis flies, or Trichoptera, of
Illinois. Bulletin Illinois Natural History Survey
23(1): 1-326.
VOLUME 99, NUMBER 2 37
bo
Sibley, C. K. 1926. Studies on Trichoptera. Prelimi- Wiggins, G. B. 1996. Larvae of the North American
nary Biological Survey of the Lloyd-Cornell Res- caddisfly genera (Trichoptera). Second edition.
ervation. Bulletin of the Lloyd Library 27: 102— University of Toronto Press, Toronto, xiii + 457
108, 185-221. pp.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 374-375
NOTE
Note on Stoneflies (Plecoptera), Particularly Prostoia besametsa (Ricker) (Nemouridae),
in the Diet of Salmonids from the Headwaters of the Middle Fork of the South Platte
River, Colorado
A previous report (Duffield and Nelson
1993. Aquatic Insects 15: 141-148) noted
that profiles of salmonid diet can provide
useful information about the relative abun-
dance and the life history of the prey items
consumed. An example of this approach
was a recent investigation of stoneflies, par-
ticularly Prostoia besametsa (Ricker)
(Nemouridae), in the diet of three Rocky
Mountain salmonid species during a limited
period of the year. Prostoia besametsa is an
abundant species, often exhibiting high
population densities in lowland creeks
throughout its range (Baumann et al. 1977.
Memoirs of the American Entomological
Society 31: 1-208). A few studies have list-
ed nemourid stoneflies in the diet of salmo-
nids (Allan 1978. Verhandlungen der inter-
nationalen Vereinigung fur Limnologie 20:
2045-2050; McNicol et al. 1985. Environ-
mental Biology of Fishes 12: 219-229;
Hubert and Rhodes 1989. Hydrobiologia
178: 225-231; Duffield and Nelson 1993),
but none reported this species.
The study area was upstream of the junc-
tion of Mosquito Creek and the Middle
Fork of the South Platte River, north of
Fairplay (Park County), Colorado (39°15'N,
and 106°04'W). A total of 38 salmonids
consisting of the brook trout (Salvelinus
fontinalis Mitchell), brown trout (Salmo
trutta L.) and cutthroat trout (Oncorhyn-
chus clarki Richardson) were sampled on
six separate days from May 29 through
June 10, 1992. The fish were collected dur-
ing the late morning and early afternoon us-
ing artificial dry flies. Stomach contents
were collected with a pump (Duffield and
Nelson 1993), placed into 4 dram vials and
preserved with 70% ethanol. Each salmonid
was measured and released unharmed.
Stoneflies in the samples were identified
to species whenever possible. Some par-
tially digested adults and immatures could
be identified only to the family or genus.
All other insects were identified to order.
Material is maintained in the authors’ col-
lections.
The lengths of the three salmonid species
sampled ranged from 160 mm to 290 mm.
A total of 2192 specimens were recovered
from 38 stomach samples, for an average of
58 insect specimens per sample. Stoneflies
were the most abundant insect order in the
diet (46% of recovered items, Table 1). All
but one sample contained at least one stone-
fly, and one sample contained eleven stone-
fly species. Five families and eleven genera
of stoneflies were identified (Table 2). All
have been reported in this drainage (Ward
and Kondratieff 1992. University Press of
Colorado, Niwot. 191 pp.).
Duffield and Nelson (1993) found that
stoneflies are important in the diet of brown
trout and rainbow trout during winter and
spring in a mid-Atlantic stream. They spec-
ulated that since many North American
stonefly species undergo nymphal diapause
in the summer and emerge from winter to
early summer, they are readily available for
Table 1. Insect orders recovered from salmonid
stomach samples taken from the Middle Fork of the
South Platte River, Colorado.
Order % of Total Specimens
Plecoptera 46.6
Diptera 22/5
Ephemeroptera 10.4
Trichoptera 9.4
Hemiptera Sa//
Hymenoptera 3)o!
Coleoptera 0.8
VOLUME 99, NUMBER 2
SS
Table 2. Plecoptera recovered from salmonid stomach samples taken from the Middle Fork of the South
Platte River, Colorado.
me aE
Nymph Adult
Family ae
Species Male Female Unknown Male Female Exuviur Total
Capniidae
Capnia confusa — — aos ats 6 ae 6
Nemouridae
Prostoia besametsa 330 548 92 + 3 2 979
Zapada haysi — — 1 —s = —_ 1
Chloroperlidae
Suwallia sp. — == 12 = = 22 12
Sweltsa sp. — = 7 — = —_ 7
Chloroperlidae sp. — == 1 — — = 1
Perlodidae
Cultus aestivalis — = 3 ae fxke a 3}
Isoperla fulva = — 1 = — ated 1
I, quinquepunctata — = 5 is bine £27: 5
Kogotus modestus a a a) —_ at 3. Lae 2
Megarcys signata — 2 1 = —— a 3
Pteronarcyidae
Pteronarcella badia — 3 a Is es = 3
Total 1023
fish consumption only during this latter pe-
riod of the year. This hypothesis is consis-
tent with stoneflies being a dominant prey
item of the Rocky Mountain salmonids ex-
amined in this study during late May and
early June.
Approximately 96% of the stoneflies re-
covered were Prostoia besametsa (Nem-
ouridae). The sample with the largest num-
ber of stonefly specimens contained 163 in-
dividuals, 161 of which were P. besametsa.
Nearly all the individuals of P. besametsa
recovered from the samples were late-instar
nymphs (Table 2). Only a few adults were
present in the samples even though they
were abundant along the river margins and
were frequently observed flying over the
river. Radford and Hartland-Rowe (1971.
Canadian Journal of Zoology 43: 1812-
1817) reported a late April to July emer-
gence for an Alberta population of P. be-
sametsa and Hassage and Stewart (1990.
Southwest Naturalist 35: 130—134) reported
a late April to May emergence for a New
Mexico population. It is possible that dur-
ing the period of this study the late-instar
nymphs of P. besametsa were intercepted
by the salmonids in the water column or on
the water surface as they were about to
emerge. The female-biased sex ratio of 1M:
1.66F late-instar nymphs indicates, as has
been reported for other stoneflies (Hynes
1976. Annual Review of Entomology 15:
25-42), that seasonal emergence of this
species is protandrous with male emergence
peaking slightly ahead of that of the fe-
males.
We thank Mark S. Schorr for his useful
comments on an earlier draft of this manu-
script.
Richard M. Duffield, Department of Bi-
ology, Howard University, Washington, DC
20059, U.S.A.; Charles H. Nelson, Depart-
ment of Biological and Environmental Sci-
ences, The University of Tennessee at Chat-
tanooga, Chattanooga, TN 37403, U.S.A.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, p. 376
NOTE
Cereal Leaf Beetle (Coleoptera: Chrysomelidae) as a Pest of Ornamental Grasses
The adventive cereal leaf beetle, Oulema
melanopus (L.) (Coleoptera: Chrysomeli-
dae), is distributed throughout Europe, ex-
tending into Siberia, northern Africa, and
Scandinavia. It was first found in the United
States in 1962 from Michigan. Since then,
it has spread throughout the eastern United
States. The cereal leaf beetle has been pres-
ent in Maryland since 1967 (Haynes and
Gage 1981. Annual Review of Entomology
26: 259-287). Since 1967, the cereal leaf
beetle has been reported only feeding on
small grains and corn.
The cereal leaf beetle is primarily a pest
of small grains, Avena sativa L. (oats), Hor-
deum vulgare L. (barley), Secale cereale L.
(rye), and Triticum aestivum L. (wheat), but
summer adults may feed on Zea mays L.
(corn). Reports in the European literature
also list several genera of Poaceae as
hosts—Agropyron repens (L.) Beauv.
(quackgrass), Avena fatua L. (wild oat),
Brachypodium pinnatum (L.) Beauv. (chalk
false broom), Dactylis glomerata L. (or-
chardgrass), Festuca gigantea (L.) Vill., H.
murinum L. (wall barley), Lolium multiflo-
rum Lam. (annual ryegrass), L. perenne L.
(perennial ryegrass), Phalaris canariensis
L. (canarygrass), and Phleum pratense L.
(timothy) (Hodson 1929. Bulletin of Ento-
mological Research 20: 5-14; Balachowsky
and Mesnil. 1953. Les Insectes Nuisibles
aux Plantes Cultivees, pp. 788-795; Bala-
chowsky. 1963. Jn Entomologie Appliqué a
Agriculture, Vol. 2; Miczulski. 1973. Ro-
czniki Nauk Rolniczych Seria E 3: 61-86;
Schmitt. 1988. pp. 475-495 In Jolivet, P,
E. Petitpierre, and T. H. Hsaio, eds., Biol-
ogy of Chrysomelidae). Most of the United
States literature on the cereal leaf beetle
deals with small grains (Battenfield et al.
1982. Bulletin of the Entomological Society
of America 28: 291-301). Wilson and
Shade (1966. Annals of the Entomological
Society of America 59: 170—173) tested the
cereal leaf beetle on a number of grasses
and found that it could feed and reproduce
on Triticum spelta L. (spelt), Phalaris arun-
dineacea L. (reed canarygrass), Bromus
inernis Leyss. (smooth brome), Elymus sp.
(wildrye), and Festuca arundineacea
Schreb. (tall fescue). Castro et al. (1965.
Quarterly Bulletin-Michigan State Univer-
sity Agricultural Experiment Station 47:
623-653) also found that the cereal leaf
beetle could reproduce on Setaria italica
(L.) Beauv. (foxtail millet).
On 9, 10, and 20 May 1996 I observed
adults and second and third instar larvae of
the cereal leaf beetle feeding on ornamental
varieties of Phalaris arundinacea (ribbon
grass) in three separate nurseries in Calvert
and St. Mary’s Counties Maryland. Also,
cereal leaf beetle adults were found feeding
on P. arundinacea in a landscaped garden
in Anne Arundel County on 2 June. Two
varieties of P. arundinacea were attacked—
‘Picta’ and ‘Strawberries and Cream’.
Feeding damage by the beetle was typical
cereal leaf beetle damage. Adjacent orna-
mental grasses in the genera Chasmanthium
(sea oats), Miscanthus, and Pennisetum
(fountain grass) were not being fed on.
Voucher specimens are deposited in the
Maryland Department of Agriculture col-
lection. Maryland Department of Agricul-
ture Contribution Number CN 96-96.
C. L. Staines, Maryland Department of
Agriculture, Plant Protection Section, 50
Harry S. Truman Parkway, Annapolis,
Maryland 21401, U.S.A.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 377-378
NOTE
Collections of Stephanidae (Hymenoptera) in the Mid-Atlantic States Including an
Eastern Record for Schlettererius cinctipes (Cresson)
The Nearctic Stephanidae includes two
genera and six species as revised by
Townes (1949. Proceedings of the United
States National Museum 99: 361-370).
They are parasitoids of wood-boring Cole-
optera and Hymenoptera. Two species are
native to eastern North America, Megischus
bicolor (Westwood) and Megischus brun-
neus Cresson, but M. brunneus occurs only
in southern Florida. Specimens of the fam-
ily are not commonly collected. In 15 years
of extensive Malaise trapping in Virginia,
Maryland, and West Virginia, I have taken
only 49 specimens. Of these, 42 were taken
in one trap at the same spot over a two-year
period in Essex Co., Virginia. All speci-
mens were M. bicolor, except for one spec-
imen of Schlettererius cinctipes (Cresson)
taken in Fairfax Co., Virginia. Schletterer-
ius cinctipes is native to the Pacific coast
states, with one record from the Black Hills
of South Dakota.
Megischus bicolor (Westwood)
This species occurs from Massachusetts
and southern Ontario south to Florida and
west to Iowa, Utah, Texas, southern Ari-
zona and southern California. The host re-
corded is “‘buprestid in Cercidium torrey-
anum [a synonym of Cercidium floridum
Benth. ex A. Gray]” (Carlson. 1979. Cat-
alog of Hymenoptera in America North of
Mexico, Vol. 1, pp. 740-741). Other rear-
ing records are emerging from Quercus, Q.
bicolor Willd., Carya, Cercidium, and Pi-
nus, and specimens have been collected
from dead Populus grandidentata Michx.
and a standing, dead Fagus grandifolia
Ehrh. (Townes 1949, Carlson 1979).
The flight of Megischus bicolor in Vir-
ginia is from the end of May into Septem-
ber, with most records in June (15), July
(16), and August (13). These records were
from traps at the edge of woods, mostly Pi-
nus spp., Quercus spp., Liquidambar styr-
aciflua L., and Liriodendron tulipifera L.
The sex ratio for trapped specimens was 22
females and 26 males.
Collection records.—MARYLAND:
Prince George’s Co., Beltsville Agricultural
Research Center, VII-20-28-93 (1 2). VIR-
GINIA: Essex Co., | mi SE Dunnsville,
V25-VESVIe Cd 6) ViEela—27.91 (G2):
VI.28—VI1I.11.91 i 2); VIT.12—26.91 (1 2;
1 do), VIL27—VIII.9.91 (1 2,1 3), VII.10—
26.9156. 2.25) Gos, V2 7X. 16:91 Cl):
VI.12—24.92 (1 9, 8 3), VI.25—VII.6.92 (3
@. 1 G);, VIL7—-17-92 (4 ¢), VILIS8—31-92
G2. 36). VITIsI— 14.92 Ol eo 2 ss
VI.10—23.93 (1 2), VII.16—VIII.1.94 (1 3),
IX.7—30.94 (1 2). Fairfax Co., nr. Annan-
dale (backyard trap), VII.16—22.93 (1 ),
VIII.14—20.96 (1 &).
Schlettererius cinctipes (Cresson)
This species is known from British Co-
lumbia, Washington, Oregon, California,
Arizona, Idaho, and South Dakota (Black
Hills). It has been reared from various co-
nifers in the West, including Abies concolor
(Gord. and Glend.) Lindl., Picea engelman-
ni Parry, Pinus ponderosa Dougl. ex Laws.,
and Pseudotsuga menziesii (Mirb.) Franco
(Townes 1949, Carlson 1979), and is a par-
asitoid of Sirex noctilio (F) (Siricidae) in
Australia where it was imported from west-
ern United States (Taylor 1967. Journal of
the Australian Entomological Society 6:
13-19; Kirk 1975. Pan-Pacific Entomolo-
gist 51: 57-61).
This is the first record from eastern Unit-
ed States. The specimen was from a trap set
in the backyard of our house in suburban
Virginia; a nearby woodpile consists mostly
378
of cut pine and maple. I have run a trap at
this spot since 1981.
Siricids are commonly transported by
commerce in wood and occasionally
emerge in building after the wood is used
for construction. Although Sirex noctilio is
not Nearctic and not in the native range of
Schlettererius cinctipes, S. cinctipes could
parasitize other Siricidae, and thus be trans-
ported in the same manner.
Collection record.—VIRGINIA: Fairfax
Co., nr. Annandale (backyard trap), VIII.4-
10.96 (1 @).
KEY TO SPECIES
The Nearctic species are keyed by
Townes (1949). The following couplet will
separate the two species collected in the
mid-Atlantic states.
1. First gastral segment about 2.5 longer than
broad (Townes 1949, fig. 1); female hindtarsus
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
5-segmented; apical half of first and entire sec-
ond and third metasomal segments orange
Moye see Sou ertome Schlettererius cinctipes (Cresson)
— First gastral segment about 4 longer than
broad (Townes 1949, fig. 2); female hindtarsus
3-segmented; metasoma black
AN to ERA crore cee Megischus bicolor (Westwood)
I thank Mr. and Mrs. John G. Kloke for
allowing field work on their property in Es-
sex Co., Virginia. The following kindly re-
viewed the manuscript: Steve L. Heydon,
University of California, Davis and Karl V.
Krombein, Smithsonian Institution, Wash-
ington, D.C.
David R. Smith, Systematic Entomology
Laboratory, PSI, Agricultural Research
Service, % National Museum of Natural
History, MRC 168, Washington, DC 20560,
U.S.A.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, p. 378
NOTE
Species Homonymy in Belomicrus Costa (Hymenoptera: Sphecidae: Crabroninae)
In 1994 two new species of the wasp ge-
nus Belomicrus Costa were described under
the name montanus. Belomicrus montanus
Kazenas and Antropov (1994. Zoologiches-
kiy Zhurnal 73: 68-77) was published in
January, and B. montanus Bohart (1994.
Journal of Hymenoptera Research 3: 207—
226) appeared November 16. Thus, Bo-
hart’s species is a junior homonym. I pro-
pose the replacement name Belomicrus bo-
harti for B. montanus Bohart. It is named
after Richard M. Bohart, in recognition of
his work on this difficult genus.
Arnold S. Menke, Ammophila Research
Institute, 1429 Franklin Street, Bisbee, AZ
85603-6211, U.S.A.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 379-398
OBITUARY
George C. Steyskal
1909—1996
George Constance Steyskal, dipterist and
former president of the Entomological So-
ciety of Washington, died in Gainesville,
Florida on May 30, 1996. He had moved to
Gainesville after the death of his wife in
1991. He is survived by a son, Neil Morgan
Steyskal of Washington, D.C., and by two
grandchildren, two great-grandchildren, and
two brothers. A talented amateur with a
publication record that few professional
taxonomists would equal, tremendous
knowledge of languages, and generous
sharer of his expertise, his career merits re-
counting.
George was born in Detroit, Michigan,
on March 30, 1909, the oldest of seven chil-
dren. Early on, he entered factory work to
help the family, graduating from the Henry
Ford Trade School in Detroit. He worked
for many years as a tool-and-die maker, fi-
nally becoming superintendent of a plant in
that craft.
Although an amateur and employed in
industry, his taxonomic papers, often ac-
companied by his own drawings, attracted
attention. When a vacancy occurred in the
U.S. Department of Agriculture’s organi-
zation for taxonomic entomology, now
known as the Systematic Entomology Lab-
oratory (SEL), he applied and was appoint-
ed, at first as an Agricultural Research
Technician, soon as Research Agriculturist.
In spite of his publications and recommen-
dations from prominent entomologists who
knew him and his abilities, his lack of for-
mal academic qualifications did not match
the established requirements for Research
Entomologist. However, his already excel-
lent publication record won him appoint-
ment to a research position, for which he
did full justice as his numerous publications
attest. He was employed at SEL in Wash-
ington, D.C., from July 2, 1962 until his
retirement on Sept. 30, 1979. He continued
to work full time at SEL and the National
Museum of Natural History, where he was
a Research Associate, until after the death
of his wife and his move to Florida, where
he became a Resident Research Associate
of the Florida State Collection of Arthro-
pods.
George had broad interests in natural his-
tory, especially in insects, snails, and bot-
any. His knowledge of plants, aided by his
phenomenal memory, was wide ranging.
Eventually he concentrated on Diptera and
especially on the Acalyptratae, to which
most of his published papers were directed.
His assignment in SEL especially included
plant-feeding Diptera of the families Te-
phritidae (fruit flies) and Agromyzidae (leaf
miners), and here his knowledge of the
plant hosts was especially valuable. No
doubt many of his papers on varied subjects
involved material that crossed his desk in
SEL for identification, but the point is that
he did something about it as well as about
new material discovered in the National
Collection at the Museum, and about lin-
guistic or nomenclatural questions brought
to him for advice.
The family Sciomyzidae was one of his
early loves, and one of his best contribu-
tions. For years, North American species of
the genus Dictya had been identified as D.
umbrarum (Linnaeus), except for five spe-
cies described by Curran in 1932. After de-
scribing a few species in 1938-1939,
George studied the genus deliberately for
some years, and in 1954 produced a defin-
itive review of the genus with eight plates
of figures of the male genitalia. By the time
George C. Steyskal at work (1977)
of the Nearctic Diptera Catalog (1965),
George had described 13 of the 19 species
then recognized in Dictya.
His publication record totals 446 titles
(See Bibliography, based on his own list
which often grouped the titles in catalogs).
The breadth and size of his knowledge, in-
terests, and contributions in Diptera are ev-
ident in the following figures:
4 new subfamilies in 4 different families;
24 new genera and 2 new subgenera in
10 families, mostly in his favorites: 9 in
Sciomyzidae, 8 in Otitidae plus Platysto-
matidae;
347 new species (including six nomina
nova and eight subspecies) in 32 families
(as presently recognized), chiefly in 22 of
Acalyptratae, with scattered species in 10
other families, all species by Steyskal alone
except for 34 co-authored with nine other
persons. All this in addition to designations
of type species and lectotypes, new synon-
ymy, corrections of authorship or dates of
publication, elevations in rank of family-
group names, and a number of pertinent
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
comments on applications in the Bulletin of
Zoological Nomenclature.
George was a good cataloger, and he nev-
er failed to respond to requests for coop-
eration in the various regional catalogs.
Thus he contributed 15 families for the Ne-
arctic Catalog (1965); later 15 Neotropical,
16 Oriental, and 5 Afrotropical. In addition,
he contributed chapters on 14 families
(three co-authored) for the Manual of Ne-
arctic Diptera (1981, 1987).
His cooperation, generosity, and ever-in-
creasing publication record were also rec-
ognized by genera and species named in his
honor. Four genera are based on his name:
Steyskalia Aczél 1959 in Micropezidae,
Laksyetsa Foote 1978 in Tephritidae, Steys-
kaliella Sods 1978 in Platystomatidae and
Steyskalomyza Kurahashi 1982 in Dryomy-
zidae. Some 28 species have been named
steyskali, but there is no up-to-date list of
these. The published regional catalogs list
ten such names but there are undoubtedly
more in the years subsequent to the cata-
logs. As a further tribute, volume 3 of the
Oriental Catalog (1977) was dedicated to
George.
Another feature of his publication record
is the number of book reviews, which often
contained pertinent comments on the au-
thors’ treatments of scientific names. Oddly
enough, in his personal list of publications
he recorded the reviews but usually did not
number them in his own total!
A remarkable facet of this amateur-be-
come-professional was his lifelong interest
in linguistics, both classical Latin and
Greek and modern languages. He translated
all (perhaps almost all!) the European lan-
guages and had studied others such as Ar-
abic and Japanese. For recreation he read
books in Italian. He was always helpful and
generous with his time as colleagues visi-
tors, and staff from around the Museum
came to him for help with a phrase, a de-
scription, a type designation, or a transla-
tion from some foreign language. This
made him a natural adviser on scientific
names for the Proceedings of the Biological
VOLUME 99, NUMBER 2
381
George identifying flies, above with Keith Harris (on right) 1981, below with Sidney Camras (on the right)
1988.
Society of Washington and the Proceedings
of the Entomological Society of Washing-
ton, which he served faithfully and prompt-
ly for over twenty years. Further, from 1980
until his death he was Translation Editor for
Scripta Tecnica’s Entomological Review
(the Russian Entomologicheskoe Obozren-
ie). In the Review for June 1996, Scripta
Tecnica published a fine tribute to his “‘un-
tiring effort, his great linguistic skills, and
his vast knowledge of entomology.”’ During
his years of service for the two Washington
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
George meeting with colleagues, above at the Montreal Congress (1956), Curtis Sabrosky, Reginald Painter,
Frank Hull and George (left to right); below at the London Congress (1964). George, Jerry Rozen and Curtis
Sabrosky (left to right).
VOLUME 99, NUMBER 2
journals, he was often able to correct errors
in names before they appeared in print, or
suggested names that better expressed the
meanings intended by authors. I can testify
personally to George’s ability as a word-
smith in devising scientific names. In his
own papers, one of his most brilliant pro-
posals was for an anomalous genus of un-
certain position whose exact place in Aca-
lyptrate Diptera was long moot. His name:
Cinderella!
His long interest in linguistics is evident
in the number of titles dealing with scien-
tific names and their correct spelling, gen-
der, stem, and similar considerations. I
counted at least 27 1945-1989, that clearly
dealt with the grammar of names, besides
some general papers and numerous items in
regular taxonomic publications. As would
be expected, most such papers concerned
Diptera, but his wide-ranging interests, and
no doubt in part the problems brought to
him for advice, resulted in papers on names
and their grammar in such varied groups as
Amphibia, Arachnida, Coleoptera, Dermap-
tera, Heteroptera, Hymenoptera, Mallopha-
ga, Plecoptera, and Psocoptera. His exam-
ples and recommendation have encouraged
many authors to include a few lines on Et-
ymology to show the derivation of their
new scientific names.
George was an excellent collector and
observer in the field, and his published pa-
pers from time to time reflected this, such
as his notes on pre-copulatory and mating
behavior in Dolichopodidae, color and col-
or pattern in the eyes of live flies, and the
dipterous fauna of tree trunks. As the years
passed, there were fewer such papers and
more on straight descriptive taxonomy.
His industry experience and natural tal-
ent were revealed many times in things he
would construct, from a large and complete
doll house for his granddaughter to a drop-
leaf table on which he and I dueled in a
couple of games of rapid chess during our
lunch breaks. He developed a method and
equipment for making non-glass microvials
for storing insect genitalia (see Gurney,
Kramer & Steyskal 1964). He was always
interested in matters of technique of all
kinds, and wrote occasional papers on such
subjects. Hence he was the logical lead au-
thor in an up-to-date revision of a large bul-
letin on “Techniques for Collection and
Preservation of Insects and Mites”’
Steyskal, Murphy & Hoover 1986).
George was a loyal member of scientific
societies, beginning with the local Detroit
Entomological Society, which evolved in
1954 into the Michigan Entomological So-
ciety, of which he served as first President
in 1955—56, and was voted Honorary Mem-
ber in 1970. He joined the Entomological
Society of Washington in 1947 (President
in 1976) and the Entomological Society of
America in 1949, continuing in both until
his death. He was also a member, at least
for some period of time, of other societies
of which we know the Biological Society
of Washington, Entomological Society of
Canada, Florida Entomological Society, and
the Society of Systematic Zoology.
Of his many interests, one that merits
special mention is his love of and abiding
devotion to classical music. He had a vast
collection of records and great knowledge
of them. He loved to sit quietly and listen
to the music, sometimes singing in Italian
along with the soloists.
Yes, George had faults, as who does not?
Those in daily contact with him, in such
matters as curation, protection of boxed
specimens from insect pests, and reaction to
manuscript committee’s comments on his
prose, were aware of some weaknesses, per-
haps not unexpected in a chiefly self-taught
individual without formal training or profes-
sorial guidance or oversight. With his brain
power, what might have he accomplished
with academic training!! Or would it have
spoiled him? His colleagues recall with
pleasure his remarkable memory, amazing
knowledge of languages, breadth of inter-
ests, and the generous helpfulness of this
multi-talented, self-taught amateur with a
record of accomplishments that eclipses that
of many professionals. As the publishers of
(see
384
Entomological Review expressed it in a trib-
ute to George in the June 1996 issue of the
Review, his professional associates “will al-
ways remember his good humor, energy,
zest for life and kindness, as well as his en-
cyclopedic learning.”’ He was truly an un-
usual individual. Having survived a quarter
century of car-pooling with George, I can
even chuckle over memories of his driving!
ACKNOWLEDGMENTS
I thank Jennifer Fairman for preparing
the plates and Chris Thompson for prepar-
ing the list of publications. The list of pub-
lications was prepared from George’s own
list, but carefully checked against the orig-
inals. George’s own annotated set of his pa-
pers is now held by the Systematic Ento-
mology Laboratory, USDA. The _ photo-
graphs were taken by Chris Thompson (fig.
2) and George Byers (figs. 3—5). These pho-
tographs and most of George’s scientific pa-
pers are now in the Smithsonian Archives.
So, finally I thank William Cox of these
Archives for his assistance.
Curtis W. Sabrosky, 205 Medford Leas,
Meadford, NJ 08055-2236, U.S.A.
SCIENTIFIC WRITINGS OF
GEORGE C. STEYSKAL
1938
New Stratiomyidae and Tetanoceridae (Diptera) from
North America. Occas. Pap. Mus. Zool. Univ.
Mich. 386, 10 pp., 2 pls. [1938.10.29 imprint date,
Steyskal marked it as received 1938.11.07]
1938
The pre-copulatory behavior of the male of Dolicho-
pus omnivagus Van Duzee (Diptera, Do-
lichopidae). Bull. Brooklyn Ent. Soc. 33: 193-
194. [1938. 11.15]
1938
Unusual pentatomid records. Bull. Brooklyn Ent. Soc.
537201 (938 ties)
1939
Notes on preparation technique. Bull. Brooklyn Ent.
Soc; 333 235, (1939/01. 13]
1939
Ennearthron oblongum. Bull. Brooklyn Ent. Soc. 34:
20. [1939.02.27]
1939
A new species of Dictya (Sciomyzidae, Diptera). Can-
ad. Ent. 71: 78. [1939.04.01]
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1939
Distributional notes on Bembicidae (Hymenoptera).
Bull. Brooklyn Ent. Soc. 34: 218-219.
[1939.09.27]
1940
A list of Michigan Diptera. Bull. Brooklyn Ent. Soc.
35: 143. [1940.11.11]
1940
& C. W. Sabrosky. Diptera of Michigan. Ent.
News 51: 242. [1940.11.08]
1940
Additional specimens of Lasia purpurata (Acro-
ceridae). Bull. Brooklyn Ent. Soc. 35: 158.
[1940.12.26]
1941
A curious habit of an Empidid fly. Bull. Brooklyn Ent.
Soc. 36: 117. [1941.06.24]
1941
A new species of Euparyphus from Michigan (Diptera,
Stratiomyidae). Bull. Brooklyn Ent. Soc. 36: 123-—
124. [1941.06.24]
1941
Odontomyia records (Diptera, Stratiomyidae). Bull.
Brooklyn Ent. Soc. 36: 125. [1941.06.24]
1941
A new species of Pterodontia (Diptera, Acroceridae).
Bull. Brooklyn Ent. Soc. 36: 140. [1941.06.24]
1942
A new species of Phyllomyza from Virginia (Diptera:
Milichiidae). Ent. News 53: 84—85. [1942.03.20]
1942
Notes on the genus Dolichopus (Diptera, Do-
lichopodidae). Paper 2. Bull. Brooklyn Ent. Soc.
37: 62-67. [1942.06.30]
1942
A curious habit of an empidid fly; further notes. Bull.
Brooklyn Ent. Soc. 37: 67 [1942.06.30]
1942
A note on preparing Diptera. Bull. Brooklyn Ent. Soc.
37: 69. [1942.06.30]
1942
Miscellaneous behavioristic observations upon Diptera
(Clusiidae, Chloropidae, Sciomyzidae). Bull.
Brooklyn Ent. Soc. 37: 101—102. [1942.08.27]
1943
Asterocampa celtis in Michigan (Lepidoptera: Nym-
phalidae). Ent. News 54: 27. [1943.02.06]
1943
A new species of Pholeomyia, with a key to the North
American species (Diptera, Milichiidae). Ent.
News 54: 99-102. [1943.04.27]
1943
Old World Sepsidae in North America, with a key to
the American genera (Diptera). Pan-Pacific Ent.
19: 93-95. [1943.07.31]
1944
A new ant-attacking fly of the genus Pseudacteon,
with a key to the females of North American spe-
VOLUME 99, NUMBER 2
cies (Diptera, Phoridae). Occas. Pap. Mus. Zool.
Univ. Mich. 489, 4 pp. [1944.12.11]
1945
Behavior of Strauzia longipennis var. vittigera Loew
(Diptera, Trypetidae). Bull. Brooklyn Ent. Soc. 39:
156. [1945.02.20]
1945
The prey of Crossocerus pammelas Pate (Sphecidae).
Bull. Brooklyn Ent. Soc. 39: 170. [1945.02.20]
1945
Note on preparing Diptera. Bull. Brooklyn Ent. Soc.
39: 170. [1945.02.20]
1945
A key to the North American species of the genus
Suillia R.-D. (Diptera, Helomyzidae). Bull.
Brooklyn Ent. Soc. 39: 173-176. [1945.02.20]
1945
Notes on Nallachius americanus (McL.) (Dilaridae,
Neuroptera). Psyche 51: 183-184. [before
1945.02.25 receipt date at USNM]
1945
Nomenclature and semantics. Ent. News 56: 100-102.
[1945.05.28 receipt date at USNM]
1945
[Anonymous] The butterflies of Wayne County, Mich-
igan. The Detroit Entomological Society, Detroit,
Michigan, [1945.06.??]
1945
Behavior of Thaumatomyia (=Chloropisca) species
(Diptera, Chloropidae). Bull. Brooklyn Ent. Soc.
40: 48. [1945.06.13]
1945
Remarks upon spatial relationships in entomological
description. Bull. Brooklyn Ent. Soc. 40: 57-59.
[1945.06.13]
1945
Leafhoppers swarming (Homoptera, Cicadellidae).
Bull. Brooklyn Ent. Soc. 40: 86. [1945.09.10]
1946
(Book Review) Foundations of Plant Geography, by
Stanley Cain. Ent. News 57: 31. [1946.04.08]
1946
The number of species in a genus. Ent. News 57: 57—
58. [1946.05.06]
1946
Themira nigricornis Meigen in North America, with a
revised key to the nearctic species of Themira
(Diptera: Sepsidae). Ent. News 57: 93-95.
(1946.06.20]
1946
[Proposal and discussion of family name Cer-
oxydidae]. P. 170-171. in C. W. Sabrosky, Family
names in the order Diptera. Proc. Ent. Soc. Wash.
48: 163-171. [1946.10.01]
1947
The genus Diacrita Gerstaecker (Diptera, Otitidae).
Bull. Brooklyn Ent. Soc. 41: 149-154.
[1947.01.30]
385
1947
The mating behavior of Tachytrechus vorax, T. moe-
chus, and Gymnopternus barbatulus (Diptera,
Dolichopodidae). Bull. Brooklyn Ent. Soc. 41:
168-169. [1947.01.30]
1947
Summa Brasiliensis Biologiae. Ent. News 58: 20.
[1947.04.17]
1947
Notes on the genus Dolichopus (Diptera, Do-
lichopodidae). Paper 3. Bull. Brooklyn Ent. Soc.
42: 34-38. [1947.05.28]
1947
Distributional notes on Lauxaniidae mostly from the
Great Smoky Mountains National Park (Diptera).
Ent. News 58: 72-73. [1947.06.13]
1947
Micropezidae (Diptera) from the Solomon Islands. Oc-
cas. Pap. Mus. Zool. Univ. Mich. 502, 10 pp., |
pl. [1947.08.04]
1947
The distribution by states of United States insect col-
lection records. Ent. News 58: 154-156.
[1947.09.30]
1947
A revision of the nearctic species of Xylomyia and Sol-
va (Diptera, Erinnidae). Pap. Mich. Acad. Sci.
Arts Letters 31: 181-190. [1947.10.??]
1947
Diostracus prasinus Loew in Tennessee (Diptera, Do-
lichopodidae). Bull. Brooklyn Ent. Soc. 42: 16.
[1947.05.28]
1947
A new genus and species of Rhopalomeridae from
northwestern Mexico (Diptera). Proc. Ent. Soc.
Wash. 49: 225-228. [1947.11.12]
1949
New Diptera from Michigan (Stratiomyidae, Sarco-
phagidae, Sciomyzidae). Pap. Mich. Acad. Sci.
Arts Letters 33: 173-180. [1949.05.02 separates
received |
1949
Sarcophagidae (Diptera) from the Great Smoky Moun-
tains National Park. Bull. Brooklyn Ent. Soc. 44:
60. [1949.04.29]
1949
Rhagionidae (Diptera) from the Great Smoky Moun-
tains National Park. Bull. Brooklyn Ent. Soc. 44:
68. [1949.04.29]
1949
A new anomalous acalyptrate fly (Diptera). Bull.
Brooklyn Ent. Soc. 44: 134-137. [1949.10.24]
1949
An indexing system for taxonomists. Coleopt. Bull. 3:
65-71. [1949.11.14 separates received]
1949
Sepsidae from the Australasian region (Diptera). Pan-
Pacific Ent. 25: 161-171. [1949.11.25]
386
1949
Notes on color and pattern of eye in Diptera. Bull.
Brooklyn Ent. Soc. 44: 163-164. [1949.12.13]
1950
The genus Protodictya Malloch (Diptera, Scio-
myzidae). Proc. Ent. Soc. Wash. 52: 33-39.
[1950.02.14]
1950
Notes and records of Phytalmiidae (Diptera: Acalyp-
tratae). Wasmann J. Biol. 8: 93—96. [1950.05.17]
1950
An easy way to make entomological drawings. Ent.
News 61: 137-139. [before 1950.10.20 receipt
date at USNM]
1950
A curious habit of an Empidid fly; third note. Bull.
Brooklyn Ent. Soc. 45: 155. [1950.12.??]
1951
The genus Sepedon Latreille in the Americas. Was-
mann J. Biol. 8: 271-297. [1951.02.15]
1951
A new species of Tetanocera from Korea [Diptera:
Sciomyzidae). Wasmann J. Biol. 9: 79-80.
[1951.05.25]
195i
The dipterous fauna of tree trunks. Pap. Mich. Acad.
Sci. Arts Letters 35: 121-134. [1951.??.??]
1951
A new species of Euparyphus from Ontario (Diptera,
Stratiomyidae). Proc. Ent. Soc. Wash. 53: 273-—
274. [1951.10.77]
1952
Insects feeding on plants of the Toxicodendron section
of the genus Rhus (Poison oak, ivy, or sumac).
Coleopt. Bull. 5: 75-77. [1952.01.??]
1952
Ulidiinae (Diptera, Otitidae) of Australasian Regions.
Occas. Pap. Bernice P. Bishop Mus. 20: 277-287.
[1952.02.22]
1952
Australasian stilt-legged flies (Diptera: Tylidae) in the
United States National Museum. Proc. U. S. Nat.
Mus. 102: 161-180. [1952.02.26]
1952
Notes and records of Phytalmiidae. II. (Diptera: Aca-
lyptratae). Wasmann J. Biol. 10: 87-90.
[1952.06.03]
1952
James, M. T. & . A review of the nearctic Stra-
tiomyini (Diptera, Stratiomyidae). Ann. Ent. Soc.
Amer. 45: 385-412. [1952.10.??]
1952
(Book Review) Taxonomy of vascular plants, by
George H. M. Lawrence. Ann. Ent. Soc. Amer.
45: 219. [1952.06.??, 1952.08.08 receipt date at
USNM]
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1952
Genus Pseudorichardia Hendel (Diptera, Otitidae).
Occas. Pap. Bernice P. Bishop Mus. 21: 63-66.
[1952.10.29]
1952
Review of ‘“Grundziige einer Theorie der phylo-
genetischen Systematik,” by Dr. Willi Hennig.
Ann. Ent. Soc. Amer. 45: 502—503. [1952.09.??,
1942.11.04 receipt date at USNM]
1953
‘“‘On the nature of systematics.”’ [Translation of a pas-
sage from Hennig’s Grundziige.] Syst. Zool. 2: 41.
[1953.77.22]
1953
Further notes on Diptera of tree trunks with descrip-
tions of two new species of Drapetis (Diptera;
Empididae) and an abstract of a Finnish paper on
Diptera by Tuomikoski. Pap. Mich. Acad. Sci.
Arts Letters 38: 255-260. [1954.??.??]
1953
Comment on ‘“‘How Many Species.”’ Evolution 7: 179.
[1954.??.??]
1953
A suggested classification of the lower brachycerous
Diptera. Ann. Ent. Soc. Amer. 46: 237-242.
[1955.08.03]
1953
A new melissomimetic fly of the genus Microdon
(Diptera, Syrphidae). Occas. Pap. Mus. Zool.
Univ. Mich. 551, 4 pp. [1956.12.31]
1954
Colobaea and Hedria, two genera of Sciomyzidae new
to America (Diptera: Acalyptratae). Canad. Ent.
86: 60—65. [1954.03.??]
1954
The genus Pteromicra Lioy (Diptera, Sciomyzidae)
with especial reference to the North American
species. Pap. Mich. Acad. Sci. Arts Letters 39:
257-269. [1954.??.??]
1954
The Sciomyzidae of Alaska (Diptera). Proc. Ent. Soc.
Wash. 56: 54-71. [1954.05.??]
1954
The American species of the genus Dictya Meigen
(Diptera, Sciomyzidae). Ann. Ent. Soc. Amer. 47:
511-539. [1954.11 .??]
1955
Notes on a collection of Argentine Sciomyzidae. Acta
Zool. Lilloana 13: 73-75. [1955.06.06 separates
received]
1956
The Eastern species of Nemomydas Curran (Diptera:
Mydaidae). Occas. Pap. Mus. Zool. Univ. Mich.
573, 5 pp., 1 pl. [1965.03.09]
1956
The genus Seioptera Kirby (Diptera, Otitidae). Ann.
Ent. Soc. Amer. 49: 30-32. [1956.03.??]
VOLUME 99, NUMBER 2
1956
New species and taxonomic notes in the family Scio-
myzidae. Pap. Mich. Acad. Sci. Arts Letters 41:
73-87. [1956.03. 26 received by section chairs]
1956
Dolichopodidae (Diptera Brachycera) from Nova Sco-
tia. Canad. Ent. 88: 139-140. [1956.05.??]
1956
& J. Verbeke. Sepedoninae (Sciomyzidae,
Diptera) from Africa and Southern Arabia. Bull.
Inst. Roy. Sci. Nat. Belgique 32 (7): 1-14. (1956.
01.??]
1957
The postabdomen of male acalyptrate Diptera. Ann.
Ent. Soc. Amer. 50: 66—73. [1957.02.??]
1957
The relative abundance of flies (Diptera) collected at
human feces. Zeitschr. Angew. Zool. 44: 79-83.
SS Ae 7]
1957
The date of publication of Bezzi’s studies in Philippine
Diptera, I. Proc. Ent. Soc. Wash. 59: 90.
W957205222)]
LOS if
A revision of the family Dryomyzidae (Diptera, Aca-
lyptratae). Pap. Mich. Acad. Sci. Arts Letters 42:
55-68. [1967.05.25 separates received]
OSH
Notes on color and pattern of eye in Diptera. II. Bull.
Brooklyn Ent. Soc. 52: 89-94. [1957.11.??]
1958
A new species of the genus Preromicra associated with
snails (Diptera, Sciomyzidae). Proc. Ent. Soc.
Wash. 59: 271-272. [1958.06.??]
1958
Notes on nearctic Helcomyzidae and Dryomyzidae
(Diptera, Acalyptratae). Pap. Mich. Acad. Sci.
Arts Letters 43: 133-143. [1958.03.31]
1958
Notes on the Richardiidae, with a review of the species
known to occur in United States (Diptera, Aca-
lyptratae). Ann. Ent. Soc. Amer. 51: 302-310.
[1958.05. 24]
1958
The genus Somatia Schiner (Diptera, Somatiidae).
Rev. Brasil. Ent. 8: 69-74. [1958.03.15]
1958
A method for the analysis of local faunal lists. Proc.
10th Internat. Congr. Ent., Montreal, !: 761-765.
[1959.12.??]
1958
Notes on North American Piophilidae (Diptera). Proc.
Ent. Soc. Wash. 60: 246. [1958.12.18]
1959
The American species of the genus Tetanocera Du-
méril (Diptera). Pap. Mich. Acad. Sci. Arts Letters
44: 55-91. [1959.03.28]
387
1959
Dolichopus correus, new species, and notes on other
Dolichopodidae (Diptera, Brachycera). Occas.
Pap. Mus. Zool. Univ. Mich. 604, 6 pp.
[1959.05.29]
1960
The genus Antichaeta Haliday, with special reference
to the American species (Diptera, Sciomyzidae).
Pap. Mich. Acad. Sci. Arts Letters 45: 17-26.
[1960.03.28 separates received]
1960
New North and Central American species of Sciomy-
zidae (Diptera, Acalyptratae). Proc. Ent. Soc.
Wash. 62: 33-43. [1960.04.27]
1960
Diptera (Brachycera) Otitidae. So. Afr. Animal Life 7:
309-315. [1960.12.??]
1961
Two new species of Sepsisoma from Kansas (Diptera,
Richardiidae). J. Kansas Ent. Soc. 34: 83-85.
[1961.05.09 separates received]
1961
The genera of Platystomatidae and Otitidae known to
occur in America north of Mexico (Diptera, Aca-
lyptratae). Ann. Ent. Soc. Amer. 54: 401-410.
[1961.05 .??]
1961
The North American Sciomyzidae related to Pherbel-
lia fuscipes (Macquart) (Diptera, Acalyptratae).
Pap. Mich. Acad. Sci. Arts Letters 46: 405-415.
[1961.06.??]
1962
The American species of the genera Melieria and
Pseudotephritis (Diptera: Otitidae). Pap. Mich.
Sci. Arts Letters 47: 247—262. [1962.05.23]
1962
The genus Curranops Harriot (Diptera: Otitidae). Proc.
Ent. Soc. Wash. 64: 117-118. [1962.08.27]
1962
Notes on palaearctic Dryomyzidae, Helcomyzidae and
Sciomyzidae (Diptera). Notulae Ent. 42: 71-72.
[1962.10.15]
1962
& R. H. Foote. Notes on the genus I/cterica
Loew in North America (Diptera: Tephritidae).
Proc. Ent. Soc. Wash. 64: 166. [1962.10.17]
1963
Dichaetophora gracilis (Loew) (Diptera, Sciomyzidae)
in England. Entomologist 96: 10. [1963.01.??]
1963
African Platystomatidae and Pyrgotidae (Diptera), in
the Carnegie Museum collections. Ann. Carnegie
Mus. 36: 133-138. [1963.02.08]
1963
A second North American species of Traginops Co-
quillett (Diptera, Odiniidae). Proc. Ent. Soc.
Wash. 65: 51—54. [1963.04.29]
388
1963
Taxonomic notes on Sciomyzidae (Diptera, Aca-
lyptratae). Pap. Mich. Acad. Sci. Arts Letters 48:
113-125. [1963.08.30 separates received]
1963
The genus Notogramma Loew (Diptera Acalyptratae,
Otitidae). Proc. Ent. Soc. Wash. 65: 195-200.
[1963.10.25]
1963
A note on Seioptera importans (Diptera: Otitidae).
Ann. Ent. Soc. Amer. 56: 878. [1963.11.??]
1963
The genus Plagiocephalus Wiedemann (Dipt. Otiti-
dae). Studia Ent. 6: 511-514. [1963.12.??]
1964
A tribe of Cecidomyiidae (Diptera) new to North
America. Proc. Ent. Soc. Wash. 66: 54.
[1964.03.05]
1964
Gurney. A. B., J. P. Kramer & . Some tech-
niques for the preparation, study, and storage in
microvials of insect genitalia. Ann. Ent. Soc.
Amer. 57: 240-242. [1964.03.19]
1964
Two new species of Platystomatidae (Diptera) from
Africa, with a key to the known species of Neoe-
pidesma Hendel. Acta Zool. Acad. Sci. Hung. 10:
263-268. [1964.03.20]
1964
Larvae of Micropezidae (Diptera), including two spe-
cies that bore in ginger roots. Ann. Ent. Soc.
Amer. 57: 292-296. [1964.05.??]
1964
Descriptive and synonymical notes on Liriomyza mun-
da (Diptera: Agromyzidae). Ann. Ent. Soc. Amer.
57: 388-389. [1964.05.??]
1964
(Book Review) British Flies. VI. Empididae. Collin, J.
E. Quart. Rev. Biol. 38: 405. [1963.12.??]
1964
(Book Review) Principles of Numerical Taxonomy.
Sokal, R. R., and P. H. A. Sneath. Ann. Ent. Soc.
Amer. 57: 390. [1964.05.??]
1964
The subgenus Neocota of the genus Rhamphomyia
(Diptera: Empididae). Ann. Ent. Soc. Amer. 57:
466—468. [1964. 07.27]
1964
Notes on North American Piophilidae II (Diptera).
Proc. Ent. Soc. Wash. 66: 177-181. [1964.10.21]
1964
The synonymy of Willineria orfilai Blanchard (Dip-
tera, Otitidae). Studia Ent. 7: 490. [1964.12.??]
1964
Two cases of abnormal development in male postab-
domens of flies of the family Sciomyzidae (Dip-
tera). Pap. Mich. Acad. Sci. Arts Letters 49: 195—
198. [1964.12.24 separates received]
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1965
A new species of the genus Xenopterella Malloch from
the United States (Diptera: Lauxaniidae). J. Kans.
Ent. Soc. 38: 66—68. [1965.02.17]
1965
Synonymy of the genera Antillonerius and Imrenerius
(Diptera: Neriidae). Proc. Ent. Soc. Wash. 67: 60.
[1965. 04.29]
1965
The genus Peocilotraphera Hendel (Diptera: Platy-
stomatidae). Proc. Ent. Soc. Wash. 67: 84-87.
(1965.07.01]
1965
The subfamilies of Sciomyzidae of the World (Diptera:
Acalyptratae). Ann. Ent. Soc. Amer. 58: 593-594.
[1965. 07.16]
1965
Families Acartophthalmidae (p. 808), Clusiidae (p.
805-807), Dryomyzidae (p. 680-681), Hel-
comyzidae (p. 678-679), Micropezidae (p. 633—
636), Neriidae (p. 637), Otitidae (p. 642-654),
Piophilidae (p. 710-713), Platystomatidae (p.
655-657), Pyrgotidae (p. 657—658), Richardiidae
(p. 641-642), Sciomyzidae (p. 685-695), Sepsi-
dae (p. 681—685), Tanypezidae (p. 641), Thyreo-
phoridae (p. 713-714). In Stone, A., C. W. Sa-
brosky, W. W. Wirth, R. H. Foote, and J. R. Coul-
son—A catalog of the Diptera of America north
of Mexico. U. S. Dept. of Agric., Agric. Res.
Serv., Agric. Handbook 276, 1696 pp.
[1965.08.23]
1965
Trend curves of the rate of species description in zo-
ology. Science 149: 880-882. [1965.08.20]
1965
The family Tanypezidae in North America (Diptera,
Acalyptratae). Proc. Ent. Soc. Wash. 67: 201.
[1965.10.06]
1965
The third larval instar and puparium of Odontoloxozus
longicornis (Diptera: Neriidae). Ann. Ent. Soc.
Amer. 58: 936-937. [1965.11.??]
1965
Notes on uninominal nomenclature. Syst. Zool. 14:
346-348. (1965.12.24]
1965
Records and descriptions of African Otitidae and Pla-
tystomatidae (Diptera). Ann. Natal Mus. 18: 171—
178. (1965.11.??]
1965
(Book Review) The families and genera of North
American Diptera, 2nd revised edition, by C. H.
Curran. Bull. Ent. Soc. Amer. 11: 102-103.
[1965.06.??]
1965
Percentage of synonymy in Diptera. Proc. Ent. Soc.
Wash. 67: 206. [1965.09.??]
VOLUME 99, NUMBER 2
1965
(letter) Convection plumes from trees. Science 150:
LGZ9O [L9G s sez
1966
A new species of Atherigona Rondani causing damage
to wheat plants in West Pakistan (Diptera: Mus-
cidae). Proc. Ent. Soc. Wash. 68: 53—56.
[1966.03.18]
1966
Notes on types of some species described in Sciomyza
and Tetanocera by Loew, Walker and Van der
Wulp (Diptera: Sciomyzidae, Muscidae, Neriidae,
Pyrgotidae). Studia Ent. 8: 445—448. (1965.02.28]
1966
Notes on the genus Pygophora Schiner, with the de-
scription of one new species (Diptera: Muscidae).
Pacific Insects 8: 14—16. [1966.05.10]
1966
The genus Eumecosomyia Hendel (Diptera, Otitidae).
Proc. Ent. Soc. Wash. 68: 100—102. [1966.06.29]
1966
A case of homonymy in the genus Hercostomus (Dol-
ichopodidae). Proc. Ent. Soc. Wash. 68: 169.
[1966.06.29]
1966
Notes on the genus Helina Robineau-Desvoidy with a
new species from Wyoming (Diptera, Muscidae).
Proc. Ent. Soc. Wash. 68: 177-179. [1966.09.20]
1966
A key to the species of the genus Xanthacrona Wulp
(Diptera, Otitidae). Proc. Ent. Soc. Wash. 68: 269.
[1966.09.20]
1966
A new species of Sciapus from California, with a re-
vised key to the species of Sciapus found in
America north of Mexico (Diptera: Dolicho-
podidae). Proc. Ent. Soc. Wash. 68: 290-294.
(1966.12.19]
1966
A review of the North American species of the genus
Otites Latreille, with descriptions of two new spe-
cies (Diptera: Otitidae). Mich. Ent. 1: 79-84.
[1966.12. 24]
1966
Otitidae from the Galapagos Islands (Diptera, Acalyp-
tratae). Proc. Calif. Acad. Sci. (4) 34: 483—497.
(page 498 blank) [1966.12.30]
1966
Notes on flies captured in treetops in Malaya (Diptera:
Empididae, Neriidae, Platystomatidae, Sepsidae,
Muscidae). Proc. U. S. Nat. Mus. 120 (3562): 1—
16. [stamped as issued 1966.12.30, but separates
not received until 1967.02.??]
1966
(Book Review) The natural history of flies, by Old-
Loyd, EH: (Quart: Revs (Biol) -41:) 21/7=2118)
[1966.06.??]
389
1967
& M. Sasakawa. Diptera: Clusiidae. Ins. Mi-
cronesia 14: 243-249. [1967.01.10]
1967
The nearctic species of Pherbellia Robineau-Desvoidy,
subgenus Oxytaenia Sack (Diptera, Sciomyzidae).
Papers Mich. Acad. Sci. Arts Letters 51: 31-38.
[1967.03.03 received Smithsonian library]
1967
A key to the species of Tritoxa Loew (Diptera: Otiti-
dae). Proc. Ent. Soc. Wash. 69: 94. [1967.03.28]
1967
A key to the genera of Anthomyiinae known to occur
in America north of Mexico, with notes on the
genus Ganperdea Aldrich (Diptera, Anthomyi-
idae). Proc. Biol. Soc. Wash. 80: 1—7. (page 8 is
blank) [1967.03.24]
1967
Notes on some older types of neotropical Micro-
pezidae with descriptions of new species (Diptera,
Acalyptratae). Papeis Avulsos Zool., S. Paulo 20:
75-83. (page 84 is blank) [1967.03.28]
1967
The generic name //ione (Diptera, Sciomyzidae). Ent.
Rec. J. Var. 79: 152-153. [1967.06.??]
1967
Palaeosepsis species described by Melander and Spu-
ler (Diptera: Sepsidae). Proc. Ent. Soc. Wash. 69:
180. [1967.08.30]
1967
Replacement names for preoccupied specific names in
Dolichopodidae (Diptera). Proc. Ent. Soc. Wash.
69: 224. [1967.10.13]
1967
Another view of the future of taxonomy. Syst. Zool.
16: 265—268. [1967.09.30]
1967
Anthomyia procellaris Rondani in North America
(Diptera, Anthomyiidae). Proc. Ent. Soc. Wash.
69: 240. [1967.10.13]
1967
A new subgenus for Leucopis luteicornis Malloch, a
predator on mealybugs in India (Diptera: Cha-
maemyiidae). Proc. Soc. Wash. 69: 275-276.
[1967.10.13]
1967
Distinguishing Amoebaleria defessa (Osten Sacken)
from A. sackeni Garrett (Diptera: Heleomyzidae).
Proc. Ent. Soc. Wash. 69: 296. (1967.10.13]
1967
Something better than polyporus or pith for double
mounts) “Broce Ent! Secs Wash? 6924297:
[1967.10.13]
1967
Family Tanypezidae. Cat. Dipt. Amer. s. U. S. 52, 4
pp. [1967.08.30]
1967
Family Pyrgotidae. Cat. Dipt. Amer. s. U. S. 56, 8 pp.
[1967.08.30]
390
1967
Family Helcomyzidae. Cat. Dipt. Amer. s. U. S. 59, 3
pp. [1967.08.30]
1967
Family Ropalomeridae. Cat. Dipt. Amer. s. U. S. 60,
7 pp. [1967.08.30]
1967
Family Coelopidae. Cat. Dipt. Amer. s. U. S. 61, 1 pp.
[1967.08.30]
1967
Family Piophilidae. Cat. Dipt. Amer. s. U. S. 68, 2 pp.
[1967.08.30]
1967
(Book Review) Dictionnaire des termes techniques
d’Entomologie élémentaire, by E. Séguy. Bull.
Ent. Soc. Amer. 13: 248. [1967.09.??]
1968
Notes on North American Piophilidae. II. (Diptera).
Proc. Ent. Soc. Wash. 70: 25—27. [1968.04.02]
1968
Priority of the generic names Tonnoiria Malloch and
Tonnoiria Parent (Diptera). Proc. Ent. Soc. Wash.
70: 87. [1968.04.02]
1968
Family Micropezidae. Cat. Dipt. Amer. s. U. S. 48, 33
pp. [1968.04.21]
1968
Family Neriidae. Cat. Dipt. Amer. s. U. S. 49, 7 pp.
[1968.04.21]
1968
Family Richardiidae. Cat. Dipt. Amer. s. U. S. 53, 26
pp. [1968.04.21]
1968
Family Otitidae. Cat. Dipt. Amer. s. U. S. 54, 31 pp.
[1968.04.21]
1968
Family Platystomatidae. Cat. Dipt. Amer. s. U. S. 55,
4 pp. [1968.04.21]
1968
Family Sepsidae. Cat. Dipt. Amer. s. U. S. 63, 6 pp.
[1968.04.21]
1968
The size factor in numerical taxonomy. Syst. Zool. 17:
210. [1968.06.28]
1968
Paratoxurinae, a new subfamily name in the Pyrgotidae
(Diptera). Proc. Ent. Soc. Wash. 70: 147.
[1968.06.29]
1968
The synonymy of Amoebaleria sackeni Garrett (Dip-
tera: Heleomyzidae). Proc. Ent. Soc. Wash. 70:
113. [1968.01.29]
1968
& Saad El-Bialy. A list of Egyptian Diptera
with a bibliography and key to families. United
Arab Republic, Ministry of Agriculture. Tech.
Bull. 3, 87 pp. [1968.09.??. While imprint date is
1967, work appeared in Sept. 1968]
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1968
The genus Somatia Schiner, If. (Diptera: Somatiidae).
Proc. Ent. Soc. Wash. 70: 255—257. [1968.10.??]
1968
Beauty vs. utility in mounting small flies on paper
points. Proc. Ent. Soc. Wash. 70: 286.
[1968.10.??]
1968
The number and kind of characters needed for signif-
icant numerical taxonomy. Syst. Zool. 17: 474—
477. [1968.12.19]
1968
A case of partially reduplicated antenna in Minettia
obscura (Loew) (Diptera: Lauxaniidae). Proc. Ent.
Soc. Wash. 70: 360. [1968.12.18]
1968
Notes and descriptions of Egyptian acalyptrate Diptera
(Ephydridae, Milichiidae, Otitidae). Bull. Soc.
Ent. Egypte 50: 109-125. [1968.12.21]
1968
(Book Review) Briefe tiber Blattminierer (Letters on
Leaf Miners), by Erich M. Hering, selected, edit-
ed, and annotated by Kenneth M. Spencer. Bull.
Ent. Soc. Amer. 14: 294. [1968.12.??]
1968
(Book Review) Trudy Vsesojuznovo Entomologi-
cheskovo Obshchestva. Bull. Ent. Soc. Amer. 14:
78. [1968.?7.??]
1969
New species of Empididae of the genera Empis, Hi-
lara, and Hormopeza from Georgia, with a syn-
opsis of the North American species of Hormo-
peza. Ann. Ent. Soc. Amer. 62: 292-299. [1969.
03.17]
1969
The postabdomen of Cinderella, with a new species
from Argentina (Diptera: Heleomyzidae). J. Kans.
Ent. Soc. 42: 80-83. [1969.04.25]
1969
The mistreatment of the Latin genitive case in forming
names of parasites. Syst. Zool. 18: 339-342.
[1969.09.??]
1970
Notes on the morphology of the male copulatory ap-
paratus in the Agromyzidae (Diptera). Proc. Ent.
Soc. Wash. 71: 531-533. [1970.01.12]
1970
The anatomy and taxonomy of the beet leaf miner of
Egypt and Cyprus, Pegomya mixta (Diptera: An-
thomyiidae). Ann. Ent. Soc. Amer. 63: 300-307.
[1970. 01.15]
1970
Distinction between Phytomyza horticola Goureau and
P. syngenesiae (Hardy) (Diptera, Agromyzidae).
Ent. News 80: 301—303. [1970.02.02]
1970
Notes on Hawaiian Otitidae (Diptera). Proc. Hawai.
Ent. Soc. 20: 439—441. [1970. 02.19]
VOLUME 99, NUMBER 2
1970
The species of the genus Dimecoenia (Diptera: Ephyd-
ridae) in America north of Panama, with the de-
scription of a new species. Ann. Ent. Soc. Amer.
63: 462-465. [1970.03.16]
1970
Revisionary notes on the genus Systenus Loew with
the description of a new species (Diptera: Doli-
chopodidae). Proc. Ent. Soc. Wash. 72: 107-111.
[1970.03.18]
1970
(Book Review) Diptera: Dolichopodidae. Appendix
(Phoridae). Joann M. Tenorio. 1960. Jn Zimmer-
mann, E. C., Insects of Hawaii, Honolulu, vol. 11,
suppl., 73 pp. Proc. Ent. Soc. Wash. 72: 136.
[1970.03.18]
1970
The language of zoological names. Syst. Zool. 19: 94—
97. [1970.03.24]
1970
On the nature and use of the suffix -ellus, -ella, -ellum
in species-group names. J. Lepidopt. Soc. 24: 38—
41. [1970.03. 26]
1970
(Book Review) The Empididae of Southern Africa
(Diptera), by Kenneth G. V. Smith. 1969. Annals
of the Natal Museum, vol. 19. Ann. Ent. Soc.
Amer. 63: 916. [1970.05.15]
1970
Ulidia omani, new species, from Israel (Diptera: Oti-
tidac) sa Procssentn SOG Washenii2 227229»
[1970.06.11]
1970
The date and grammar of Loxosceles, the name of a
genus of venomous spiders. Proc. Ent. Soc. Wash.
72: 280 [1970.06.11]
1970
(Book Review) The Flies of Western North America,
E R. Cole. Mich. Ent. 3: 31. [1970.06.22]
1970
On gender concord in binomina. Coleopt. Bull. 24: 57—
58. [1970.06.25]
1970
Notes on the postabdomen and variation in Centrion-
cus prodiopsis Speiser (Diptera: Diopsidae). Proc.
Ent. Soc. Wash. 72 (3): 325-327. [1970.09.25]
1970
The date of publication of Folia Entomologica Hun-
garica, vol. III. Proc. Ent. Soc. Wash. 72: 339.
[1970.09.25]
1970
On the Diptera described as Tipula fasciata by early
authors. Proc. Ent. Soc. Wash. 72 (3): 413.
[1970.09.25]
1970
Family Somatiidae. Cat. Dipt. Amer. s. U. S. 94, 2 pp.
[1970.09.10]
1970
A new North American species of Tetanops Fallén
sensu stricto (Diptera, Ottitidae). J. Kans. Ent.
Soc. 43: 378-380. [1970.11.13]
1970
Dolichopodidae from the Patuxent Wildlife Refuge,
Maryland, with the description of three new spe-
cies of Neurigona (Diptera). Proc. Ent. Soc. Wash.
72: 448-453. [1970.12.14]
1971
[Anonymous] New Western Hemisphere Record —An
anthomylid fly (Emmesomyia socia (Fallén)) —
New York, Coop. Econ. Ins. Rpt. 21: 5. [1971.
01.01]
197]
Notes on the genus Strongylophthalmyia Heller, with
a revised key to the species (Diptera: Stron-
gylophthalmyiidae). Ann. Ent. Soc. Amer. 64:
141-144. [1971.01.??]
1971
Ensina sonchi (Linnaeus) in South America (Diptera:
Tephritidae). J. Wash. Acad. Sci. 60: 158-159.
[1971.01.??]
1971
The family position and additional descriptive data
concerning Tenuia nigripes Malloch (Diptera:
Pseudopomyzidae). Pacific Insects 12: 871-873.
[1971.03. 08, imprint date incorrect, see statement
on last page of issue]
1971
(Book Review) Le Genre Rhyacophila et la Famille
des Rhyacophilidae (Trichoptera), by E Schmid.
1970. Bull. Ent. Soc. Amer. 17: 70. [1971 .??.?2]
1971
The species related to Minettia obscura (Loew), with
one new species and one new synonym (Diptera:
Lauxaniidae). Proc. Ent. Soc. Wash. 73: 17—22.
[1971.03.22]
1971
The subgenus Tritolestes Ghesquiére of the genus
Cryptochetum Rondani with a new species from
Pakistan (Diptera: Cryptochetidae). Proc. Ent.
Soc. Wash. 73: 48-51. [1971.03.22]
1971
On the validity of Cicindela cyanocephalonota Eck-
hoff (Coleoptera: Cicindelidae). Coleopt. Bull. 25:
34. [1971.03. 28]
1971
The genus Trigonosoma Gray (=Tropidogastrella
Hendel) (Diptera: Platystomidae). J. Wash. Acad.
Sci. 61: 26-28. [1971.04.26]
1971
Sepsis costalis Wiedemann redescribed and referred to
Richardiidae (Diptera, Acalyptratae). Papeis Avul-
sos Zool., S. Paulo 23: 157—160. [1971.02.12]
1971
A second species of the genus Jdanophana Hering
(Diptera, Otitidae). Papeis Avulsos Zool., S. Paulo
25: 31-33. [1971.03.31]
592
1971
Notes on the grammar of acarine nomenclature. Acar-
ologia 12: 639-642. [1971.??.??]
1971
The genus Paraleucopis Malloch (Diptera: Chamae-
myiidae), with one new species. Ent. News 82: 1—
4. [1971.05.??]
1971
A note on Camptoprosopella equatorialis Shewell
(Diptera: Lauxaniidae). Proc. Ent. Soc. Wash. 73:
110. [1971.06.28]
1971
Spilochroa geminata Sabrosky a synonym of S. polita
(Malloch) (Diptera: Trixoscelididae). Proc. Ent.
Soc. Wash. 73: 157. [1971.07.28]
1971
On the grammar of names formed with-scelus, -sceles,
-scelis, etc. Proc. Biol. Soc. Wash. 84: 7-11. (page
12 is blank) [1971.06.30]
1971
The case of Psoidos, Psodos or Psolos Treitschke. Z.N.
(SS) 362""Bull Zools Nomencle 28>) Lie
[1971.08.10]
1971
A new Central American species of Zacompsia Co-
quillett, with a key to the described species (Dip-
tera: Otitidae). Proc. Ent. Soc. Wash. 73: 247-248.
[1971.10.06]
1971
The genus Calobatina Enderlein (Diptera: Micropezi-
dae). Proc. Ent. Soc. Wash. 73: 327-328.
[1971.10.06]
1971
(Book Review) Aquatic Diptera (1934-7), O. A. Jo-
hannsen; Part V by L. C. Thomsen. Los Angeles:
Entomological Reprint Specialists, 1969. Mich.
Ente selves omlelle 23)
1971
Notes on the genera Homalomitra Borgmeier, Pycno-
pota Bezzi, and Sphinctomyia Borgmeier (Dip-
tera: Sphaeroceridae). Proc. Ent. Soc. Wash. 73:
376-378. [1971.12.17]
1971
A note on the nominate character of Paracantha den-
tata Aczél (Diptera: Tephritidae). Proc. Ent. Soc.
Wash 73: 398. [1971.12.17]
1971
Delphinia picta (Fabricius) in Central America (Dip-
tera: Otitidae). Proc. Ent. Soc. Wash. 73: 445.
[1971.12.17]
1971
Notes on some species of the genus Copromyza, sub-
genus Borborillus. J. Kans. Ent. Soc. 44: 476—
479. [1971.12.30]
1971
On the grammar of the name Heliothis Ochsenheimer
(Noctuidae). J. Lepidopt. Soc. 25: 264-266.
(LST AR IE ?2))
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1972
The taxonomy of the alfalfa blotch-miner, Agromyza
frontella (Rondani). Coop. Econ. Ins. Rept. 22:
134-137. [1972. 03.17]
1972
An unusual habit for a fly of the family Ephydridae.
Proc. Ent. Soc. Wash. 74: 129. [1972.03.28]
1972
A correction to Sturtevant’s paper on the seminal re-
ceptacles, etc., of the Diptera, 1926. Proc. Ent.
Soc. Wash. 74: 130. [1972.03.28]
1972
Euaresta reticulata (Hendel), new combination (Dip-
tera: Tephritidae). Proc. Ent. Soc. Wash. 74: 130.
[1972.03.28]
1972
The dipterous fauna of the heads of Bidens pilosa re-
examined. Florida Ent. 55: 87—88. [1971.06.01]
972
(Book Review) The Anthomyiidae of California exclu-
sive of the subfamily Scatophaginae (Diptera). By
H. C. Huckett, 1971, etc. Proc. Ent. Soc. Wash.
74: 132. [1972.03.28]
1972
Comment on the gender of Leiopelma Fitzinger, 1861
(Amphibia). Z.N.(S.) 1936. Bull. Zool. Nomencl.
295 2-a 9722 05,0)i
1972
Application for correction of certain names on the Of-
ficial List of Family-Group Names in Zoology.
Z.N.(S.) 1965. Bull. Zool. Nomencl. 29: 26—27.
[1972.05.01]
1972
A catalogue of species and key to the genera of the
family Diopsidae (Diptera: Acalyptratae). Stutt-
gart. Beitr. Naturk. 234, 20 pp. [1972.08.??, im-
print date of 1972.02.01 wrong, proof read in
May, separates received in August]
1972
New and little-known Agromyzidae from Michigan
(Diptera: Acalyptratae). Great Lakes Ent. 5: 1-10.
[1972.08.18]
1972
The genus Acrometopia Schiner in North America
(Diptera: Chamaemyiidae). Proc. Ent. Soc. Wash.
74: 302. [1972.10.26]
1972
Notes on the genus Suwillia in Mexico, with the descrip-
tion of a new species (Diptera, Heleomyzidae).
Proc. Ent. Soc. Wash. 74: 303-305. [1972.10.26]
1972
A new species of Myoleja with a key to North Amer-
ican species (Diptera: Tephritidae). Florida Ent.
55: 207-211. [1972.09.11]
L972
African Pyrgotidae (Diptera) collected by Willi Rich-
ter, Erwin Lindner, and others. Stuttgart. Beitr. Na-
turk. 238, 11 pp. [1972.07.20]
VOLUME 99, NUMBER 2
1972
Two new species of Melanagromyza Hendel (Diptera,
Agromyzidae) that bore in tomato stalks in Co-
lombia and Ecuador. J. Wash. Acad. Sci. 62: 265—
267. [1972.11.22]
1972
Dichetophora (Neosepedon) boyesi Steyskal, new spe-
cies. Pp. 362-363. Jn Boyes, J. W., Knutson, L.
V. & van Brink, J. M., Further cytotaxonomic
studies of Sciomyzidae, with description of a new
species, Dichetophora boyesi Steyskal (Diptera,
Acalyptratae). Genetica 43: 362-363. [1972.??.??]
1972
A preliminary key to the species of the neotropical
genus Tetreuaresta Hendel (Diptera: Tephritidae).
Proc. Ent. Soc. Wash. 74: 403-405. [1972.12.30]
1972
A preliminary key to the species of Neotephritis Hen-
del (Diptera: Tephritidae). Proc. Ent. Soc. Wash.
74: 414-416. [1972.12.30]
1972
Phytoliriomyza montana Frick, new synonym of P.
arctica (Lundbeck) (Diptera: Agromyzidae). Proc.
Ent. Soc. Wash. 74: 471. [1972.12.30]
1972
Expansion of Article 11 (g) (i) (4) of the Code to in-
clude provision for adjectival genitives, the form
of which is the same as the nominative. Z.N.(S.)
1969. Bull. Zool. Nomencl. 29: 135-136.
STZ See?)
1973
The meaning of the term ‘sibling species’. Syst. Zool.
(1972) 21: 446. [1973.01. 16]
1973
Pteromicra inermis Steyskal a synonym of Sciomyza
varia (Coquillett). Proc. Ent. Soc. Wash. 75: 83.
[1973.04.26]
1973
Notes on the grammar of names in the Dermaptera.
Papis Avulsos Zool., S. Paulo 26: 253-257.
[1973.02.26]
1973
A new species of the genus Archiborborus Duda from
Mexico (Diptera: Sphaeroceridae). J. Kans. Ent.
Soc. 46: 154-157. [1973.04.30]
1973
A leafminer of ragweeds (Ambrosia spp.), Calycomyza
ambrosiae (Frick) (Diptera, Agromyzidae). Coop.
Econ. Ins. Rpt. 23: 329-330. [1973.06.04]
1973
The genus Emmesomyia Malloch in North America
(Diptera, Anthomyiidae). Coop. Econ. Ins. Rpt.
23: 331-332. [1973.06.04]
1973
The genus Axiologina Hendel (Diptera: Otitidae). Flor-
ida Ent. 56: 132—134. [1973.06.27]
1973
Notes on the growth of taxonomic knowledge of the
Psocoptera and on the grammar of the nomencla-
393
ture of the order. Proc. Ent. Soc. Wash. 75: 160—
164. [1973.07.27]
1973
The identity of Calycomyza jucunda (Wulp) (Diptera,
Agromyzidae). Proc. Ent. Soc. Wash. 75: 191—
194. [1973.07.10]
1973
Dolichopus reticulus Van Duzee a synonym of D. oc-
cidentalis Aldrich (Diptera: Dolichopodidae).
Proc. Ent. Soc. Wash. 75: 239. [1973.07.10]
1973
On the family-group names based upon the genera
Bucculatrix Zeller and Stenoma Zeller (Lepidop-
tera). Proc. Ent. Soc. Wash. 75: 247. [1973.07.10]
1973
The genera of the family Thyreophoridae and the spe-
cies of the genus Omomyia, with one new species
(Diptera). Ann. Ent. Soc. Amer. 66: 849-852.
[1973. 07.16]
1973
The North American species of Dolichopus Latreille,
group B (Diptera, Dolichopodidae). J. Kans. Ent.
Soc. 46: 347-359. [1973.07.31]
1973
Distinguishing characters of the walnut husk maggots
of the genus Rhagoletis (Diptera, Tephritidae).
Coop. Econ. Ins. Rept. 23: 522. [1973.08.03]
1973
The genus Euphya Wulp, with a new species from the
Philippines (Diptera: Pyrgotidae). Pacific Insects
15: 209-212. [1973.07.20 imprint date, but re-
ceived in Washington, D.C., [1973. 09.10]
1973
A new classification of the Sepedon group of the fam-
ily Sciomyzidae (Diptera) with two new genera.
Ent. News 84: 143-146. [1973.09.17]
1973
Distinguishing larvae of the two American species of
Musca (Diptera, Muscidae). Coop. Econ. Ins. Rpt.
23: 677-678. [1973.09.28]
1973
Euxesta penacamposi Steyskal, new species. P. 226. In
Campos S., L., & L. E. Pena G., Los insectos de
Isla de Pascua, (Resultados de una prospecci6n
entomoldgica). Rev. Chil. Ent. 7: 217-229.
[1973.27.22]
1973
The grammar of names in Slater’s Catalogue of the
Lygaeidae of the World (Heteroptera). Proc. Ent.
Soc. Wash. 75: 276-279 [1973.10.05]
1973
A further note on Acompha costalis (Wiedemann)
(Diptera: Richardiidae). Proc. Ent. Soc. Wash. 75:
S25 | MOS MOOS)
1973
A term strictly equivalent to ““Type-species.”’ Proc.
Ent. Soc. Wash. 75: 377. [1973.10.05]
394
1973
The strange fate of the “‘serpentine leaf miner” (Lir-
iomyza spp., Agromyzidae, Diptera). Coop. Econ.
Ins. Rpt. 23: 735-736. [1973.10.26]
1973
, H. Robinson, H. Ulrich & R. L. Hurley. Hy-
drophorus Fallén, 1823 (Insecta, Diptera, Doli-
chopodidae): Request for suppression under the
plenary powers of the designation by Macquart,
1827 of H. jaculus Fallén as type of the genus in
favour of H. nebulosus Fallén in order to conserve
consistent usage. Z.N.(S.) 2036. Bull. Zool. No-
mencl. 30: 118—120. [1973.??.??]
1973
The grammar of names in the catalogue of the Miridae
(Heteroptera) of the world by Carvalho, 1957—
1960. Studia Ent. 16: 203—208. [1973.10.31]
1973
The Clusiidae (Diptera) of Puerto Rico and the Virgin
Islands. Studia Ent. 16: 439—444. [1973.10.31]
1974
Notes on southeastern North American species of Con-
dylostylus and Sciapus (Diptera: Dolichopodidae).
Florida Ent. 56: 359-360. 1974.02.05]
1974
The genus Dictyodes Malloch (Diptera: Sciomyzidae).
Proc. Ent. Soc. Wash. 75: 427—430. [1974.02.12]
1974
Additional data on Tephritis unicolor Walker, 1837,
newly referred to the genus Lamproxynella Hering
(Diptera: Tephritidae). Proc. Ent. Soc. Wash. 76:
49-51. [1974.04.30]
1974
Euxesta mazorca, new species, associated with ears of
maize in South America (Diptera, Otitidae). Proc.
Biol. Soc. Wash. 87: 73-75. (page 76 is blank)
[1974.04.25]
1974
A gynandromorphic specimen of the genus Limnia
(Diptera: Sciomyzidae). J. Wash. Acad. Sci. 64:
11. [1974.27.22]
1974
Sabrosky, C. W. & . The genus Sobarocephala
(Diptera: Clusiidae) in America north of Mexico.
Ann. Ent. Soc. Amer. 67: 371-385. [1974.05.15]
1974
Recent advances in the primary classification of the
Diptera. Ann. Ent. Soc. Amer. 67: 513-517.
[1974.05.15]
1974
The genus Oedopa Loew (Diptera, Otitidae Ulidiinae).
Proc. Ent. Soc. Wash. 76: 163-164. [1974.07.22]
1974
A new species of Procecidochares (Diptera: Tephriti-
dae) causing galls on stems of Hamakua Pamakani
(Ageratina riparia: Asteraceae) in Hawaii. Coop.
Econ. Ins. Rept. 24: 639-641. [1974.08.09]
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1974
Further comment on the valid emendation of -ii end-
ings of patronyms. Z.N.(S.) 1913. Bull. Zool. No-
mencl. 30: 137. [1974.06.28]
1974
& D. K. McAlpine. Platystomatidae Schiner,
1862: proposed conservation as a family-group
name over Achiidae Fleming, 1821 (Insecta, Dip-
tera), Z.N.(S.): 2053. Bull. Zool. Nomencl. 31:
59-61. [1974.07.31]
1974
(Book Review) Agromyzidae (Diptera) of Economic
Importance, by Kenneth A. Spencer. Bull. Ent.
Soc. Amer. 20: 212. [1974.06.17]
1974
Proponent’s comments on effect of Article 29(d) on
the application for correction of certain names on
the Official List of Family-Group Names. Z.N.
(S.) 1965. Bull. Zool. Nomenl. 31: 113-114.
[1974.09.20]
1974
The species of the genus Euthyplatystoma Hendel
(Diptera, Platystomatidae). Proc. Ent. Soc. Wash.
76: 310-311. [1974.11.07]
1974
A new species of Zonosemata Benjamin from Colom-
bia (Diptera: Tephritidae). J. Wash. Acad. Sci. 64:
234-235. [1974.10.14]
1974
The distribution by states of United States insect re-
cords. Bull. Ent. Soc. Amer. 20: 283-284.
[1974.12.77]
1974
(Translation) On the life and influence of J. W. Meigen
by J. A. Forster, Stettin. Entomol. Ztg. 1846. 67—
74; 130-140. Mosquito Systematics 6: 79-88.
[1974. ??.22]
1975
—& L. V. Knutson. The cochleate vesicle, a highly
specialized device for sperm transfer in male scio-
myzid flies. Ann. Ent. Soc. Amer. 68: 367-370.
[1975. 03.??]
1975
Recognition characters for larvae of the genus Zono-
semata (Diptera, Tephritidae). Coop. Econ. Ins.
Rpt. 25: 231-232. [1975.03.21]
1975
Genus Callopistromyia Hendel (Diptera, Otitidae).
Ent. News 85: 147-148. [1975.04.18]
1975
Note on the use of the suffixes -fer and -ger in nomencla-
ture. Proc. Ent. Soc. Wash. 77: 58. [1975.04.29]
1975
The family-group names based on the name of the ge-
nus Elmis Latreille (Coleoptera). Proc. Ent. Soc.
Wash. 77: 59-60. [1975.04.29]
NOS
The gender of the genus-name Stomoxys Geoffroy,
VOLUME 99, NUMBER 2
1762 (Diptera, Muscidae). Proc. Ent. Soc. Wash.
77: 163. [1975.04.29]
1975
Anastrepha obliqua (Macquart) the prior name for
Anastrepha mombinpraeoptans Sein (Fruit Flies,
Tephritidae, Diptera). Coop. Econ. Ins. Rpt. 25:
357-358. [1975.05.02]
SHS
Comment on the request for a ruling on the stem of
family-group names based on the type genus Pet-
romyzon Z.N.(S.) 2045. 3. Bull. Zool. Nomencl.
S22 ea 97 5:03:27)
1975
(Book Review) Delfinado, M. D., and D. E. Hardy,
eds. 1973. A catalog of the Diptera of the Oriental
Region. Volume 1, Suborder Nematocera. Hono-
lulu, The University Press of Hawaii. Atoll Re-
search Bull. 185: 28. [1975.01.??]
1975
The North American species of Piophila subgenus AI-
lopiophila with black forecoxa (Diptera: Piophil-
idae). Proc. Ent. Soc. Wash. 77: 212-213. [1975.
07.23]
1975
Comment on the proposal to conserve Hydrophorus
Fallén, 1823 and to suppress the designation of
Macquart of Hydrophorus iaculua Fallén as type-
species. Z.N.(S.) 2036. Bull. Zool. Nomencl. 32:
79-80. [1975.06.27]
1975
& L. V. Knutson. Key to the genera of Scio-
myzidae (Diptera) from the Americas south of the
United States, with descriptions of two new gen-
era. Proc. Ent. Soc. Wash. 77: 274-277. [1975.
10.09]
1975
The author of the scientific name of the cabbage mag-
got. Coop. Econ. Ins. Rept. 25: 910. [1975.12.19]
1975
A new series of Enlinia from Oklahoma (Diptera: Dol-
ichopodidae). Proc. Ent. Soc. Wash. 77: 478—480.
[1975.12.23]
1975
(Book Review) Taschenlexikon zur Biologie der In-
sekten, by W. Jacobs and M. Renner. Bull. Ent.
Soc. Amer. 21: 249. [1975.12.15]
1976
Notes on the nomenclature and taxonomic growth of
the Plecoptera. Proc. Biol. Soc. Wash. 88: 408—
410. [1976.??.??]
1976
The terminology of bristles on the upper back of the
head in the higher Diptera. J. Kans. Ent. Soc. 49:
155-159. [1976.04.26]
1976
(Book Review) Pacific Insects. Monograph 31; Mono-
graph 32; by D. Elmo Hardy, etc. Quart. Rev.
Biol. 51: 324-325. [1976.06.??]
1976
Additional data on holly leafminer adults (Diptera,
Agromyzidae: Phytomyza ilicis-ilicicola group).
Coop. Plant Pest Rept., APHIS, USDA 1 (42):
767-770. [1976.10.22]
1976
A new species of leafmining fly on mealycup sage
(Diptera: Agromyzidae). Coop. Plant Pest Rept. 1:
820-822. [1976.10.29]
1976
Knutson, L., , J. Zuska & J. Abercrombie. Fam-
ily Sciomyzidae. Cat. Dipt. Amer. s. U. S. 74, 24
pp. [1976.12.08]
1976
Nicolson, D. H. & . The masculine gender of
the generic name Styrax Linnaeus (Styracaceae).
Taxon 25: 581-587. [1976.11.??]
7/7
Two new neotropical fruitflies of the genus Anastre-
pha, with notes on generic synonymy (Diptera,
Tephritidae). Proc. Ent. Soc. Wash. 79: 75-81.
[1977.01. 24]
1977
& R. H. Foote. Revisionary notes on North
American Tephritidae (Diptera), with keys and de-
scriptions of new species. Proc. Ent. Soc. Wash.
79: 146-155. [1977.01.24]
1977
History and use of the McPhail trap. Florida Ent. 60:
11-16. [1977.??.??]
1977
Pictorial key to species of the genus Anastrepha (Dip-
tera: Tephritidae). Ent. Wash., 35 pp.
[1977.09.??]
1977
Beet leafminer and spinach leafminer now considered
Soc.
to be one species Pegomya hyoscyami (Panzer)
(Diptera, Anthomyiidae). Coop. Plant Pest Rept.
2: 818. [1977.10.14]
1977
Richardia eburneosignata Hennig (Diptera: Richardi-
idae), a new record for the genus north of Mexico.
Proc. Ent. Soc. Wash 79: 648. [1977.10.26]
1977
Correction of species-names dedicated to Dr. S. En-
drody-Younga. Folia Ent. Hung. 29: 153.
[1976.05.31]
1977
Families Cypselosomatidae (p. 5—6), Neriidae (p. 8—
11), Micropezidae (p. 12-20), Strongyloph-
thalmyiidae (p. 21—23), Megamerinidae (p. 28—
29), Nothybidae (p. 30—31), Diopsidae (p. 32-36),
Pyrgotidae (p. 37—43), Platystomatidae (p. 135—
164), Otitidae (p. 165-167), Dryomyzidae (p.
173), Piophilidae (p. 229), Chyromyidae (p. 240),
Odiniidae (p. 242) Cryptochetidae (p. 396-397).
In Delfinado, M. D. & Hardy, D. E. (eds.), A Cat-
396
alog of the Diptera of the Oriental Region, Vol. 3,
x + 854 pp. [1977.27.27]
1977
& M. Sasakawa. Family Tethinidae. P. 394—
395. In Delfinado, M. D. & Hardy, D. E. (eds.),
A Catalog of the Diptera of the Oriental Region,
Vol. 3, x + 854 pp. [1977.??.??]
1977
Family Dryomyzidae. Cat. Dipt. Amer. s. U. S. 62, 2
pp. [1977.11.23]
1978
What is systematic entomology? Proc. Ent. Soc. Wash.
80: 43-50. [1978.01.31]
1978
A new pest of chili peppers in Colombia (Diptera:
Lonchaeidae). Coop. Plant Pest Rept. 3: 72.
[LOT 8e7 2277)
1978
Synopsis of the North American Pyrgotidae (Diptera).
Proc. Ent. Soc. Wash. 80: 149-155. [1978.05.04]
1978
, T. W. Fisher, L. Knutson & R. E. Orth. Tax-
onomy of North American flies of the genus Lim-
nia (Diptera, Sciomyzidae). Univ. Calif. Publ. Ent.
83: 1-48, pls. 1-5. [1978.07.??]
1978
& K. A. Spencer. A new species of Phytoli-
riomyza Hendel feeding on Jacaranda in Califor-
nia (Diptera: Agromyzidae). Coop. Plant Pest
Rept. 3: 583-586. [1978.10.13]
1978
A new species of Hylemya from Wyoming (Diptera:
Anthomyiidae). Proc. Ent. Soc. Wash. 80: 553—
555. [1978.10.20]
1978
(Remarks (p. 171) in discussion appended to) L. A.
Mound. 1978. The availability of insect collec-
tions for taxonomic research. Jn Kim, K. C., ed.
The changing nature of entomological collections:
Uses, functions, growth and management. Ent.
Scand. 9: 146-177. [1978.07.01]
1978
Helosciomyzinae in Australia (Diptera: Sciomyzidae).
Austr. J. Zool. 26: 727-743. [1978.12.27]
1979
Miscellaneous comments. [on proposed revision of In-
ternational Code of Zoological Nomenclature].
Bull. Zool. Nomencl. 35: 138-144. [1979.02.01]
1979
The gender of substantivated adjectives; with a pro-
posal for the simplification of Articles 30.a.i (2)
Z.N.(S.) 2259. Bull. Zool. Nomencl. 35: 168-174.
[1979.02.01]
1979
Reference of Cephalia fulvicornis and Cephalia ma-
culipennis (Diptera: Otitidae) to the genera Eux-
esta (Diptera: Otitidae) and Rivellia (Diptera: Pla-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tystomatidae). Proc. Ent. Soc. Wash. 81: 255-256.
[1979.06.26]
1979
The Gender of Nosopon Hopkins (Mallophaga). Proc.
Ent. Soc. Wash. 81: 325. [1979.06.26]
1979
Lutzomyia vexator (Coquillett) the correct designation
for Lutzomyia vexatrix of authors (Diptera: Psy-
chodidae). Proc. Ent. Soc. Wash. 81: 423.
[1979.08.13]
1979
Biological, anatomical, and distributional notes on
the genus Callopistromyia Hendel (Diptera:
Otitidae). Proc. Ent. Soc. Wash. 81: 450—455.
[1979.08.13]
1979
Taxonomic studies on fruit flies of the genus Urophora
(Diptera, Tephritidae). Ent. Soc. Wash., 61 pp.
[1979.10.??]
1980
Haplopeodes, a new genus for Haplomyza of authors
(Diptera, Agromyzidae). Proc. Ent. Soc. Wash. 82:
140-151. [1980.01.24]
1980
Knutson. L., Hodges, R. W., & . Format for
reporting identifications. Bull. Ent. Soc. Amer. 26:
27-28. [1980.03.15]
1980
Two-winged flies of the genus Dasiops (Diptera: Lon-
chaeidae) attacking flowers or fruit of species of
Passiflora (passion fruit, granadilla, curuba, etc.).
Proc. Ent. Soc. Wash. 82: 166—170. [1980.04.18]
1980
Orth. R., & T. W. Fisher. A new species of
Pherbellia Robineau-Desvoidy with notes on the
P. ventralis group (Diptera: Sciomyzidae). Proc.
Ent. Soc. Wash. 82: 284—292. [1980.04.18]
1980
(Descriptions of Sepedon noteoi and S. oriens). Pp.
117-120. In Hardy, D. E., & M. D. Delfinado,
Insects of Hawaii, vol. 13 —Diptera: Cyclorrhapa
III. University of Hawaii, Honolulu. [1980. ??.??]
1980
Mathis, W. N. & —. A revision of the genus Oedo-
parena Curran (Diptera: Dryomyzidae: Dryomy-
zinae). Proc. Ent. Soc. Wash. 82: 349-359.
[1980.07.11]
1980
The species of the genus Suillia found in the Americas |
south of the United States (Diptera: Heleomyzi-
dae). Proc. Ent. Soc. Wash. 82: 401—404. [1980.
07.11]
1980
The grammar of family-group names as exemplified
by those of fishes. Proc. Biol. Soc. Wash. 93: 168—
177. [1980.07.09]
1980
Errors in nomenclatural grammar in Krombein et al.,
VOLUME 99, NUMBER 2
Catalog of Hymenoptera in America North of
Mexico (1979). Sphecos 3: 43—46. [1980.??.??]
1980
Distributional and synonymical notes on Hylemya (Pe-
gohylemyia) species. Proc. Ent. Soc. Wash. 82:
553-555. [1980. 10.24]
1980
Pyrgotidae (p. 556-562), Platystomatidae (p. 563—
574), Otitidae (p. 575-577), Neriidae (p. 578), Mi-
cropezidae (p. 579-582). In Crosskey, R. W., et
al., eds., Catalogue of the Diptera of the Afrotrop-
ical Region. British Museum (Natural History).
London, 1437 pp. [1980.??.??]
1980
(Book Review) Gotch, A. E, Mammals —their Latin
names explained; a guide to animal classification.
J. Mammal. 61: 581-583. [1980.08.??]
1980
A plain-letter romanization for Russian. Libr. Re-
sources and Techn. Serv. 24: 170-173.
[1980.??.??]
1981
Six new North American species of Melanagromyza
Hendel (Diptera, Agromyzidae). J. Wash. Acad.
Sci. 70: 36—43. [1981.02.16]
1981
Orth, R. E., & . A new species of Pherbellia
Robineau-Desvoidy separated from a previously
described North American species (Diptera: Scio-
myzidae). Proc. Ent. Soc. Wash. 83: 99-104.
[1981.02.17]
1981
& Knutson, L. V. 1981. Empididae. Pp. 603—
624. In McAlpine, J. FE, et al., Manual of Nearctic
Diptera, Vol. 1. Research Branch Agriculture Can-
ada Monograph 27. vi, 674 p. [1981.??.??]
1981
(Book Review) The insects and arachnids of Canada.
Part 1. Collecting, preparing, and preserving in-
sects, mites, and spiders. Compiled by J.E.H. Mar-
tin. Proc. Ent. Soc. Wash. 83: 181. [1981.02.17]
1981
A bibliography of the Malaise trap. Proc. Ent. Soc.
Wash. 83: 225-229. [1981. 04.30]
1981
A new “bobo” fly from the Gulf of California (Dip-
tera: Chamaemylidae: Paraleucopis mexicana).
Proc. Ent. Soc. Wash. 83: 403—405. [1981.08.19]
1981
A new species of Rhagoletotrypeta (Diptera: Tephrit-
idae) from Texas, with a key to the known species.
Proc. Ent. Soc. Wash. 83: 707-712. [1981.10.28]
1981
(Book Review) Taxonomists’ Glossary of Mosquito
Anatomy. By R. E. Harbach and K. L. Knight.
Proc. Ent. Soc. Wash. 83: 805—809. [1981.10.28]
1982
A second species of Ceratitis (Diptera: Tephritidae)
397
adventive in the New World. Proc. Ent. Soc.
Wash. 84: 165-166. [1982.02.16]
1982
McAlpine, J. EF & . A revision of Neosilba
McAlpine with a key to the world genera of Lon-
chaeidae (Diptera). Canad. Ent. 112: 105-137.
[1982. 02.??]
1982
An Oriental fly found in Trinidad (Diptera: Platysto-
matidae). Proc. Ent. Soc. Wash. 84: 859-860.
[1982.10.29]
1982
A key to the genera of the subfamily Otitinae of the
Americas south of the United States (Diptera: Oti-
tidae). Mem. Ent. Soc. Wash. 10: 139-144.
[1982.09.30]
1983
(Book Review) Sunflower Species of the United
States. By C. E. Rogers, T. E. Thompson, and G.
Ip SSilee lPioes lil, Soe» WSN. ss ~ 7/0:
[1983.11.09]
1983
Further gender corrections—Catalog of Hymenoptera
in America North of Mexico. Sphecos 6: 22—23.
[1983.??.??]
1984
(Book Review) Anopheline Names: Their Derivation
and History. James B. Kitzmiller. Bull. Ent. Soc.
Amer. 30: 77. [before 1982.11.09 receipt date at
USNM]
1984
A linguistic look at the stage-stadium-instar problem.
Ann. Ent. Soc. Amer. 77(2): iti. [before
1984.04.20 receipt date at USNM]
1984
A synoptic revision of the genus Aciurina Curran,
1932 (Diptera, Tephritidae). Proc. Ent. Soc. Wash.
86: 582-598. [1984.07.17]
1984
Lectotype designation for Rhamphomyia abdita Co-
quillett (Diptera: Empididae). Proc. Ent. Soc.
Wash. 86: 668. [1984.07.17]
1984
(Book Review) Check List of the Lepidoptera of
America North of Mexico 1983. Edited by Ronald
W. Hodges and others. Proc. Ent. Soc. Wash 86:
728-729. [1984.07.17]
1985
(Letter to Editor) From type to holotype. Bull. Ent.
Soc. Amer. 31(3): 2. [1985.??.??]
1985
(Book review) Revision of the genus Archytas (Dip-
tera: Tachinidae) for America north of Mexico: F
William Ravlin and Frederick W. Stehr. Florida
Ent. 67: 580. [1985.01.11]
1985
(Book review) Butterflies East of the Great Plains. By
398
Paul A. Opler and George O. Krizek. Proc. Ent.
Soc. Wash. 87: 257. [1985.01.31]
1985
(Letter to Editor) Place of Systematics in Society. Bull.
Ent. Soc. Amer. 31(1): 6. [before 1985.04.19 re-
ceipt date at USNM]
1985
(Letter to Editor). The same language. Bioscience 35:
266—267. [1985.??.??]
1986
A new species of Taeniaptera from the United States
(Diptera: Micropezidae). Proc. Ent. Soc. Wash.
88: 174-178 [1986.01.15]
1986
Spencer, K. A. & . Manual of the Agromyzidae
(Diptera) of the United States. U. S. Dept. Agr.,
Agr. Res. Serv., Agr. Handbook 638, vi + 478 p.
[1986.05. ??]
1986
(Book review) A taxonomic treatise on the Dacidae
(Tephritoidea, Diptera) of Africa, H. K. Munro.
Bull. Ent. Soc. Amer. 32: 114. [1986.??.??]
1986
Griffiths,G.C.D., K. A. Spencer & . Napomyza
Westwood, 1840 (Insecta, Diptera): Proposed con-
servation by the suppression of Napomyza Curtis,
1837. Z. N. (S.) 2495. Bull. Zool. Nomencl. 43:
170-172. [1986.07.09]
1986
Spencer, K. A. & . Agromyza Fallén, 1810 (In-
secta, Diptera). Proposed validation of Agromyza
reptans Fallén, 1823 as type species. Z. N. (S.)
2395. Bull. Zool. Nomencl. 43: 183-185.
[1986.07.09]
1986
Taxonomy of the adults of the genus Strauzia Robi-
neau-Desvoidy (Diptera, Tephritidae). Insecta
Mundi 1: 101-117. [1986.??.??]
1986
, W. L. Murphy & E. M. Hoover, editors. In-
sects and Mites: Techniques for Collection and
Preservation. U. S. Dept. Agr., Agr. Res. Serv.,
Misc. Publ. 1443, [v] + 103 pp. [1986.12.??]
1987
(Book Review) An English Translation of Russian
Common names of Agricultural and Forest Insects
and Mites, by A. Yusha, M. E. Martignonk, and
P. Iwai. Bull. Ent. Soc. Amer. 33: 45. [1987.??.??]
1987
A new species and synonymy in the Ulidiinae (Dip-
tera, Otitidae). Florida Ent. 70: 187-188.
[1987.03.27]
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1987
Families Micropezidae (p. 761—767), Neriidae (p.
769-771), Tanypezidae (p. 773-776), Strongylo-
phthalmyiidae (p. 777-779), Psilidae (p. 781-—
784), Otitidae (p. 799-808), Platystomatidae (p.
809-812), Pyrgotidae (p. 813-816), Tephritidae
(with R. H. Foote) (p. 817-829), Richardiidae (p.
833-837), Dryomyzidae (p. 923-926), Ropalom-
eridae (p. 941-944), Sepsidae (p. 945-950). In
McAlpine, J. F, et al., Manual of Nearctic Diptera,
Vol. 2, Research Branch Agriculture Canada,
Monograph No. 28: p. i-vi, 675-1332
[1987.03.31]
1988
On the naming of higher taxa. Syst. Zool. (1987) 36:
400. [1988.03.10]
1988
Languages of entomology. Bull. Ent. Soc. Amer. 34:
53-54. [before 1988.06.23 receipt date at USNM]
1988
(Letter to Editor). How big the list, catalogue, data-
bases? Antenna 12: 38—39. [1988.??.??]
1989
More on the grammar of species names in -fer and -
ger. Sphecos 18: 4—5.
1989
(Book Review). Catalog of the Heteroptera, etc., by T.
L. Henry and R. C. Froeschner; and Noctuidae,
by R. W. Poole. Proc. Ent. Soc. Wash. 91: 649-—
650. [1989.10.05]
1990
(Book Review) The Torre-Bueno Glossary of Ento-
mology, compiled by Stephen W. Nichols, etc.
Proc. Ent. Soc. Wash. 92: 173-175. [1990.01.30]
1990
A note on hagiobouctony. Taxon 39: 243. [1990.05.??]
1990
Notes on the Trapherinae (Diptera: Platystomatidae).
Ann. Soc. Ent. Fr. (N. S.) 26: 437-441.
[1990.??.??]
1991
On the meaning of the term ‘trichobothrium.’ Ent.
News 102: 95-96. [before 1991.06.21 receipt date
at USNM]
1991
Descriptions of new species of Otitidae from Chile
(Diptera). Nuevas especies de Otitidae de Chile
(Diptera). Acta Ent. Chilena 16: 25-28.
[1991.04.??]
1995
& K. Ahlmark. Two new species of Euxesta
Loew (Diptera: Otitidae). Insecta Mundi 9: 189—
193, [1995.12577)
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 399-401
BooK REVIEW
Riceland Spiders of South and Southeast
Asia. By A. T. Barrion and J. A. Litsinger.
CAB International, Wallingford, Oxon,
United Kingdom. 1995. 700 pp., 16 color
plates Paperback. ISBN 0 85918 967 5.
$225.00.
Perhaps one of the main differences be-
tween taxonomy and any other scientific
discipline is the fact that in taxonomy, at
least so far, bad work can live almost eter-
nally. If one were to publish some truly ab-
surd and erroneous scientific article, in al-
most any scientific discipline, the work
would be immediately ignored by the sci-
entific community and the contribution
would quickly disappear into oblivion. Req-
uiescat in pace. This is not true in taxono-
my. One can describe and name new taxa
in the worst possible way you can imagine,
and as far as the publication criteria set
forth by the International Code of Zoolog-
ical Nomenclature are met, those names and
descriptions will be “‘valid’’ ones and the
scientific community will have to live with
it. To a large extent quality may not matter;
taxonomists cannot ignore a publication just
because, let us say, the particulars of the
description make any reliable identification
virtually impossible. Subsequent genera-
tions of taxonomists will have to live with
that name, no matter how bad (or good) the
original description was, to the extent that
a good fraction of their work will have to
be devoted to correcting earlier mistakes. In
this sense I predict that ““Riceland spiders
of South and Southeast Asia’ (RSSSA)
will be with us for a long, long time.
RSSSA treats more than three hundred
species of spiders collected over a period of
13 years in rice ecosystems in the Philip-
pines. Species collected in other five loca-
tions in South and Southeast Asia are also
treated in this book. According to the ab-
stract, the 342 species covered represent
131 genera and 26 spider families. A large
fraction of the species treated (257 species
or 75%) are reported as “‘new species.”’ In
addition, eight new genera are described,
seven of them are monotypic.
The book is divided in thirteen sections.
In the first two sections, Barrion and Litsin-
ger offer a historical background of ara-
neology in the Philippines and introduce the
basics of spider external anatomy. The fol-
lowing four sections treat life history (it ac-
tually deals almost exclusively with eggsacs
and first instars), materials and methods,
and the classification and key to the spider
families (in reality it only covers the spider
families that have been found in rice eco-
systems in South and Southeast Asia). The
bulk of the book, more than 600 pages, is
in the seventh section which is concerned
with the description of all the taxa. The next
four sections provide a brief description of
some other arachnid orders, some com-
ments on the diversity of spiders in Phil-
ippine rice ecosystems, the distribution
maps of riceland spiders, and a glossary of
arachnological terms. The book ends with
the bibliography section and a species in-
dex. The book is profusely illustrated with
aesthetically pleasing pen and ink drawings
by Danilo Amalin and a few color photo-
graphs.
Dalingwater (1996. Newsletter of the
British arachnological Society. 76: 6—7) has
made extensive comments on the poor ed-
itorial quality of this book. I will not ad-
dress those aspects here but instead will fo-
cus on the quality of the taxonomic work
of RSSSA.
Barrion and Litsinger have succeeded in
providing us with a real compendium. A
compendium of what not to do in taxono-
my. I will comment here on only a few of
these points. For example, I have not found
a single reference to examination of type
specimens, except those of the new taxa de-
scribed in this book (incidentally, Barrion
400
and Litsinger do not tell us where the types
of the 257 presumably new species they
have described are deposited). Is it possible
to discover so many ‘“‘new taxa” without
consulting the types of the many species
that have already been described in that part
of the world? I do not think so, precisely
for the reasons I have brought up in the first
paragraph. Barrion and Litsinger describe
eight new genera (seven of them are mono-
typic) but in most cases these genera are
known from only one or two specimens.
Since this is not a revision of any of the
groups treated I do not see any compelling
reasons for describing a new genus from a
single female specimen (e.g. Gambaque-
zonia). In fact, I can think of numerous rea-
sons for not doing so. The lack of diagnosis
for many of these new genera (e.g. Kakai-
banoides or Landopo) adds another prob-
lem to it. Some of the family names and
higher level delimitations used in the book
are out of date. For example, Nephilinae is
no longer an araneid subfamily, but a te-
tragnathid subfamily. Comaroma is not a
theridiid, but a member of the Anapidae.
New species are frequently described with-
out a diagnosis and in many cases are based
on a single specimen. At least one of the
species names (the oonopid Oopaea batan-
guena) is an incorrect original spelling (Ar-
ticle 27 of the ICZN explicitly forbids the
use of diacritic marks in scientific names)
and will have to be corrected. The theridiid
Theridion kambalum, known only from the
female holotype, is described, presumably
for the first time, without providing an il-
lustration of the epigynum.
The authors state in the Introduction that
the goal of their book is to provide a guide
that can be used by specialists and novices
to identify Philippine spiders. This is a very
laudable goal, but I wonder about the need
of describing so many new taxa in a book
that is intended mainly as an identification
guide. The book is based on spider collec-
tions in rice fields that span for more than
12 years, but, despite their efforts, the ma-
jority of the new species treated in the book
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
are known from only one or a very few
specimens (perhaps because they are not
common in rice ecosystems). Given the title
of the book, it may have been then advis-
able to treat only those species that are
commonly found in rice fields. This would
certainly make the task of identifying spi-
ders form rice fields easier for some poten-
tial users of such a guide, like IPM spe-
cialists or agricultural entomologists.
I cannot evaluate in detail the taxonomy
of all the families treated in this book, but
I would like to address the quality of the
taxonomic work of Barrion and Litsinger
by making a few remarks on the Tetrag-
nathidae. Judging from their original de-
scription, the spider they describe as Phon-
ognatha guanga N. sp. is not a Phonogna-
tha, not even a tetragnathid, but a linyphiid
of the genus Neriene. Meta baywanga and
M. tiniktirika are also misplaced since they
also are linyphiids (possibly species of Ner-
iene), not tetragnathids. All these taxa are
treated under the family Metidae, a group-
ing abandoned by most taxonomists over
the last decade. The Nephilinae have also
been recognized as a group within Tetrag-
nathidae for more than a decade, but we see
them treated here as araneids. Of the six
different species of Nephila that the authors
claim that exist in the neotropics, only two,
Nephila clavipes and N. sexpunctata, were
recognized by Levi and von Eicksted in
their revision of 1989. These examples are
not minor taxonomic changes hidden in ob-
scure publications, but widely accepted
changes that can be easily found, for ex-
ample, in Platnick’s spider taxonomy cata-
logs (N. I. Platnick 1989. Advances in Spi-
der Taxonomy 1981-1987, Manchester
University Press; 1993. Advances in Spider
Taxonomy 1988-1991, New York Ento-
mological Society and The American Mu-
seum of Natural History). What transpires
from the treatment of Tetragnathidae is a
number of errors that are too large to ig-
nore: inadequate coverage of the taxonomic
literature, gross misplacement of species in
higher taxonomic categories, genitalic illus-
VOLUME 99, NUMBER 2
trations not detailed enough to allow accu-
rate identifications, no allusion to exami-
nation of type material, lack of diagnosis in
descriptions of new taxa, no evidence of
peer review, etc.
It is obvious that much time, resources,
and dedication went into the production of
RSSSA. I have no doubts that the authors
tried to accomplish their job as best as they
could, but the final product certainly leaves
much to be desired. This book also leaves
many open questions. Was this manuscript
ever reviewed by specialists before it was
accepted for publication? If not, what kind
of scientific policies at the agency respon-
sible for the publication of RSSSA allow
something like this to happen? Could this
happen again? Could RSSSA have been
better produced as a true identification
guide for a wider audience, not as an at-
tempt to deal with the taxonomy of Phil-
ippine spiders?
401
The high price alone will keep this book
mainly confined to institutional libraries.
Based on the quality of the product it is
difficult to recommend the purchase of
RSSSA, but then again, quality is not a
guiding criterion for the taxonomists when
it comes to use the available literature. Sys-
tematists working with the spider fauna of
southern Asia will have to use this refer-
ence given the enormous, quantitatively
speaking, nomenclatural weight of the
book.
I would like to thank Jonathan Codding-
ton for his comments on a draft of this re-
view.
Gustavo Hormiga, Department of Bio-
logical Sciences, George Washington Uni-
versity, Washington, D.C., 20052 and De-
partment of Entomology, NHB -105, Smith-
sonian Institution, Washington, D.C.,
20560.
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 402-403
REPORTS OF OFFICERS
EDITOR
Volume 98 of the Proceedings totaled
844 pages. Eighty-one articles, eight notes,
three book reviews, one obituary, the By-
laws of the Society, membership list of the
Society, and instructions for authors were
published.
In addition, Memoir 17 “Contributions
on Hymenoptera and Associated Insects,
Dedicated to Karl V. Krombein”’ edited by
B. B. Norden and A. S. Menke was pub-
lished in 1996. Memoir 18, ““Contributions
to Diptera,” a volume in memory of W. W.
Wirth, edited by W. N. Mathis and W. L.
Grogan, will be published in December.
Memoir 19, ““Monograph of the Berytidae,
or Stilt Bugs (Heteroptera) of the Western
Hemisphere”’ by T. J. Henry, was approved
by the Executive Committee and will ap-
pear in 1997. Approval was also given by
the Executive Committee to print a second
edition of ‘A Handbook of the Families of
Nearctic Chalcidoidea (Hymenoptera),”’ by
E. E. Grissell and M. E. Schauff; this will
appear in 1997.
I extend my appreciation to Gary L. Mil-
ler for his continuing efforts to acquire in-
formative book reviews, to Tom Henry and
Wayne Mathis for their work on memoirs
and kindly offering advice, and to Marie
Blair for her assistance in handling corre-
spondence and routing manuscripts. With-
out their help, my job would have been
much more difficult.
I am also grateful to the many reviewers
for their time-consuming efforts and con-
structive reviews. Their contributions are
essential to help increase the quality of pa-
pers published in the Proceedings.
David R. Smith, Editor
PRESIDENT-ELECT
The 1996 Annual Banquet was held on
15 May 1996 at the Associates Court of the
National Museum of Natural History in
Washington, D.C. 93 tickets were sold (12
to students). Members and guests of the En-
tomological Society of Washington were
joined by members of the Maryland Ento-
mological Society. The Master of Ceremo-
nies for the evening was Dr. David Furth.
The speaker was Dr. May Berenbaum of the
University of Illinois at Champagne-Urbana
who spoke on “Images of the Entomolo-
gist—Moving or Otherwise.”
Receipts from the sale of tickets were
$1914.00, however the actual cost of the
event was $1997.45 (DAKA) and 372.75
(SI Audio-Visuals) for a total of $2370.20.
The difference of $456.20 was subsidized
by the society and included complimentary
dinner for the speaker and the Master of
Ceremonies. In addition, the Executive
Committee authorized a $500.00 honorari-
um for the speaker. Dr. Ray Gagné was the
most outstanding ticket salesperson selling
50 tickets.
M. Alma Solis, President-Elect
VOLUME 99, NUMBER 2 403
SUMMARY FINANCIAL STATEMENT FOR 1996
Special
General Publications Total
Fund Fund Assets
Assets: November 1, 1995 $713 145.72 $109,977.60 StL 12332
Total Receipts for 1996 TH8A7T 38 ZIPS Ol 99 11 SS5
Total Disbursements for 1996 70,065.45 — 23,678.92— 93,744.37
Assets: October 31, 1996 8,897.65 107,678.92 116,497.50
Net Changes in Funds Sie 51293 S23 77-75) S. 9.374.118
Audited by the Auditing Committee, December 3, 1996, consisting of Norman E. Wood-
ley, Thomas J. Henry, and FE Christian Thompson, Chairman. Presented to the membership
at the meeting of December 5, 1996.
Respectfully submitted,
Michael G. Pogue, Treasurer
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 404-407
SOCIETY MEETINGS
1013th Regular Meeting—October 3, 1996
The 1013th Regular Meeting was called
to order at 8:01 pm by President Ralph Eck-
erlin on 3 October 1996 in the Waldo
Schmitt Room, National Museum of Natu-
ral History, Washington, D.C. Nine mem-
bers and six guests were present. The min-
utes of the May 2 and May 15 meetings
were read by Acting Recording Secretary
Gabriela Chavarria, and approved as read.
President Eckerlin called for office re-
ports. David Smith made comments on the
status of the Memoirs.
Program Chair David Furth announced
that the speaker for the November Regular
Meeting would be Christopher Marshall
from the Department of Entomology, Cor-
nell University.
President Eckerlin received a letter from
the Washington Academy of Sciences, af-
filiated with the National Academy of Sci-
ences; Chris Thompson is our representa-
tive or delegate to the Washington Acade-
my of Sciences and they will keep us in-
formed of upcoming events.
No new membership applications were
received.
Ralph Eckerlin called for any Old or New
Business. There was none and he requested
that anyone with notes or exhibitions come
forward. Ed Saugstad brought a newspaper
with some humorous insect news. President
Eckerlin brought a monograph of the World
Bank who has committed 80 million dollars
for integrative pest management.
Dr. Thomas K. Wood was the speaker for
the evening. He is a professor at the De-
partment of Entomology and Applied Ecol-
ogy, University of Delaware. His presenta-
tion was entitled ““Speciation of Enchenopa
Treehoppers: Past and Future Studies.’’ He
kindly submitted the following abstract for
publication in the Society minutes:
Evidence for a particular hypothesis of
speciation is most often inferential and
rarely amenable to direct experimental
tests. He discussed the establishment and
the initial results associated with a long-
term experimental test of a hypothesized
speciation mechanism. The hypothesis
tested was that shifts or changes in host
plant use can initiate a process that leads
to specialization and divergence. Specif-
ically, the experiment was designed to
examine the initial events associated with
specialization through host shifts when
gene flow is not precluded. For experi-
mental purposes he selected a member of
the Enchenopa binotata species complex
(Homoptera: Membracidae) associated
with the host plant Viburnum.
To simulate two alternative modes of
host specialization, seven environments
using large walk-in outdoor cages were
created, and replicated four times. Each
cage contains two large plants at opposite
ends, separated by approximately 8.0 m.
Different combinations of the host plants
in the genus Viburnum were tried. The
first four represent allopatric environ-
ments, while the remaining three repre-
sent sympatric environments, where gene
flow among insects associated with each
host species is not precluded.
Female treehoppers were painted and
then released on each plant in the seven
environments. Oviposition was moni-
tored. Deposition of over 12,000 egg
masses was directly observed during the
first year. Since the release plant for each
female within cage is known, the tenden-
cy of females to disperse from their re-
lease plant can be determined, and gene
flow of insects between plants in the first
generation of a host shift can be estimat-
ed. The timing of egg hatch, survival, de-
velopment time, eclosion, and gene flow
during mating and oviposition were de-
termined. These results showed the role
of host fidelity or philopatry in the initial
stages of a host shift.
VOLUME 99, NUMBER 2
Six visitors were introduced. Lin Chung-
Ping, Michael Adams and Ding Liu from
the University of Delaware, Astrid Caldas,
Research Associate in Entomology, and
Alejandro Lopera from Colombia working
with Dr. Adrian Forsyth, Research Asso-
ciate in Entomology.
The meeting was adjourned at 9:16 pm.
Refreshments were kindly provided by
Ralph Eckerlin.
Respectfully submitted,
Gabriela Chavarria for Darlene Judd,
Recording Secretary
1014th Regular Meeting—November 7,
1996
The 1014th Regular Meeting was called
to order at 8:04 pm by President Ralph Eck-
erlin on 7 November 1996 in the Waldo
Schmitt Room, National Museum of Natu-
ral History, Washington, D.C. Sixteen
members and 14 guests were present. The
minutes of the October 3rd meeting were
read by Acting Recording Secretary Dawn
Southard, and approved as read.
President Eckerlin called for officer re-
ports. Alma Solis gave the report of the
President-Elect which provided details of
the 1996 Annual Banquet.
Three new member applications were re-
ceived and their names read: Charles L.
Brockhouse, Fernando Munoz-Quesada,
and Steve W. Chordes III.
David Smith reported that the Bylaws of
the Society were published in the July issue
and he provided copies for anyone interest-
ed along with copies of the Instructions for
Authors.
Program Chair David Furth announced
that the speaker for the December Regular
Meeting would be Astrid Caldas, a Re-
search Associate from the Department of
Entomology, National Museum of Natural
History, who will be speaking on the ecol-
ogy and evolution of dead leaf butterflies.
The Slate of Officers for 1997 was read
by the Chair of the Nomination Committee,
Wayne Mathis, who also expressed thanks
405
to the officers of last year for their outstand-
ing performance. The Slate of Officers
which will be voted on during the Decem-
ber 5th Regular Meeting include: M. Alma
Solis, President; Warren E. Steiner, Presi-
dent-Elect; Harold Harlan, Recording Sec-
retary; Hollis B. Williams, Corresponding
Secretary; Gary L. Miller, Custodian; Mi-
chael G. Pogue, Treasurer; David G. Furth,
Program Chair; David Adamski, Member-
ship Chair; David R. Smith, Editor; and
Ralph P. Eckerlin, Past President. The other
members of the nominating committee,
Thomas J. Henry and EF Christian Thomp-
son, were thanked for their participation.
Wayne Mathis also announced that Mem-
oir 18 in honor of Willis W. Wirth is due
this year and is awaiting the second page
proof from Allen Press. It was suggested
that David Furth could use some money to
supplement honoraria or other expenses.
Ralph Eckerlin reported that during the
Executive Committee Meeting issues that
needed to be addressed were the allotment
of space for back issues, and monetary con-
cerns.
Ralph Eckerlin asked if there were any
new members and none were present. He
called for any Old or New Business. There
was none and he requested that anyone with
notes or exhibitions come forward. Warren
E. Steiner remarked on the passing of Ted
Spilman on the 22nd of September, 1996,
and that there is a committee being formed
to prepare his obituary. Ted Spilman was
the main historian for the Society over the
last several years, and he presented a his-
tory of the Society at the 1000th Meeting
of the Society. Ted Spilman served as for-
mer Treasurer and President of the Society.
Ralph Eckerlin requested a moment of si-
lence in his memory. Ed Saugstad brought
a sample of entomological T-shirts for dis-
play, and Joyce Utmar also had one for dis-
play. David Furth presented a joke aimed
at molecular systematists.
Joyce Utmar, a life member of the Soci-
ety, presented a poem entitled “‘Beetles”
written by Ted Spilman. She also presented
406
her “‘specimen,’’ Mark Grant, that she re-
ported represented a new genus and species,
Grantius marcus.
David Furth introduced the speaker for
the evening, Chris Marshall, who is a grad-
uate student at Cornell University in the
Department of Entomology. His presenta-
tion was entitled, ‘“‘Passalidae from A to Z:
The Biology and Systematics of the Social
Beetle Family.”’ The aim of his talk was to
serve as an introduction to the remarkable
scarabaeoid family, Passalidae. First, the
characteristics of all passalids as well as
their phylogenetic position among other
scarab families was discussed. He presented
various results from studies based on mor-
phology, life history, social behavior, and
communication. The role of passalids in
ecological systems as decomposers and cre-
ator of microhabitats by way of their gal-
leries was discussed. This talk stressed the
critical role of phylogenetic research both
in general and specifically with reference to
passalid biology. Examples of the utility of
phylogenetic information in passalid biolo-
gy, such as biogeography, conservation, ad-
aptation and coevolutionary processes,
were presented. He concluded his talk by
discussing his current research on the genus
Verres (tribe: Proculini) and his pians to an-
swer questions related to brachyptery and
coevolution between these beetles and the
mites found associated with them. Upon
conclusion of his talk, Chris Marshall
passed around live specimens of passalids
from Georgia.
Four visitors were introduced: Wenhua
Lu from Rhode Island, working on mor-
delid beetles; Jean-Michel Maes, Museo
Entomologia, Nicaragua; Diomedes Quin-
tero, University of Panama, working on
scorpions and mutilids; and Dr. Volker-
Hollmann-Schirrmacher, arranging for a
Postdoctoral Fellowship in the Department
of Entomology which will begin in January.
The meeting was adjourned at 9:38 pm.
Refreshments were kindly provided by Da-
vid Furth and Gabriella Chavarria.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Respectfully submitted,
Dawn Southard for Darlene Judd,
Recording Secretary
1015th Regular Meeting—December 5,
1996
The 1015th Regular Meeting was called
to order at 8:04 pm by President Ralph Eck-
erlin on 5 December 1996 in the Waldo
Schmitt Room, National Museum of Natu-
ral History, Washington D.C. Thirteen
members and 11 guests were present. The
minutes of the November 7th meeting were
read by Acting Recording Secretary Dawn
Southard. A correction to the minutes was
made by Wayne Mathis. This correction
stated that Norman E. Woodley was a mem-
ber of the nominating committee not E
Christian Thompson. The records were later
checked and indeed the minutes were cor-
rect as read and FE Christian Thompson was
on the nominating committee not Norman
E. Woodley.
President Eckerlin called for office re-
ports. Dawn Southard read the report of the
Membership Chairman (David Adamski).
President Ralph Eckerlin read the Corre-
sponding Secretary’s Report (Hollis B. Wil-
liams). The Treasurer’s Report was read by
M. Alma Solis and the Editor’s Report was
read by David Smith.
The outgoing President Ralph Eckerlin
gave thanks to his executive committee
members which include the Recording and
Corresponding Secretaries, the Editor and
his aids, the Treasurer, and the Program
Chair. President Eckerlin pointed out that
these are the hardest working members and
whom the Society’s survival has depended
on. The President also expressed having en-
joyed his tenure serving as the Society’s
President.
President Eckerlin turned the attention of
the meeting to the Election of New Offi-
cers. The Chair of the Nominations Com-
mittee, Wayne Mathis, read the Slate of Of-
ficers and noted that two of the Officers are
not voted on, that of the current President-
Elect and the Past President. The Slate of
VOLUME 99, NUMBER 2
Officers include: M. Alma Solis, the Pres-
ident-Elect now becomes the President in
accordance with the Bylaws; Warren E.
Steiner, Jr., President-Elect; Harold Harlan,
Recording Secretary; Hollis B. Williams,
Corresponding Secretary; Gary L. Miller,
Custodian; Michael G. Pogue, Treasurer;
David G. Furth, Program Chair; David
Adamski, Membership Chair; David R.
Smith, Editor; and Ralph Eckerlin, Past
President. President Ralph Eckerlin opened
the floor for any further nominations; none
were offered. Wayne Mathis responded that
we should have a vote of the membership
on the Slate of Officers. All present were in
favor and there was no opposition.
President Eckerlin asked if there was any
Old or New Business. M. Alma Solis in-
quired about the possibility of changing the
date of the January meeting to the 9th be-
cause so many people would be away due
to the holidays. Program Chair David Furth
responded that the speaker was flexible and
could be here for either the 2nd or the 9th
of January. The President put the matter on
the floor for a vote. Twenty two people vot-
ed in favor to change the date to the 9th
and two opposed. The January meeting will
be scheduled for the 9th.
President Eckerlin requested that anyone
with notes or exhibits come forward. David
Furth brought the newly published 3 vol-
ume set on Chrysomelid Biology which is
on the market for $450.00.
David Furth announced that the speaker
407
for the January meeting would be Charles
Bellamy, a curator of Coleoptera at the
Transvaal Museum in South Africa. The
proposed title of the talk is “Beetle Col-
lecting in Southern Africa (An Inordinate
Fondness: Habitats, and Habitus).’’ David
Furth introduced our speaker for the eve-
ning, Astrid Caldas, originally from Brazil,
and now here in the Department of Ento-
mology. The presentation, entitled “‘A
Lively View of Dead-Leaf Butterflies,’ was
about the population structure, mortality
factors, and levels of mortality in imma-
tures of the neotropical dead-leaf butterfly
Anaea ryphea. The talk also looked at the
intraspecific variation in Anaea ryphea
adults and in a related species, Anaea eu-
rypyle, and the possible taxonomic impli-
cations.
Four visitors were introduced: Pamela
Burns, USDA-APHIS out of Gulfport, Mis-
sissippi; Manual Balcazar working on his
Ph.D. at the University of Florida in
Gainesville; Luis Jolly from the Universi-
dad de Venezuela; and Cathy Anderson,
museum technician with the Systematic En-
tomology Laboratory.
Past President Ralph Eckerlin presented
the gavel to President M. Alma Solis who
wished everyone a Happy Holiday, and the
meeting was adjourned at 9:18 pm. Re-
freshments were kindly provided by Ralph
Eckerlin.
Respectfully Submitted,
Dawn Southard for Darlene Judd,
Recording Secretary
PROC. ENTOMOL. SOC. WASH.
99(2), 1997, p. 408
RECENT AND UPCOMING PUBLICATIONS OF THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON
Memoir No. 17
Contributions on Hymenoptera and Associated Insects,
Dedicated to Karl V. Krombein. Edited by B. B. Norden and A. S. Menke.
216 pp. 1996 (cost: $25.00).
A Festschrift in recognition of Dr. Karl V. Krombein, Smithsonian Institution,
Washington, D.C. A collection of 22 invited papers, mostly on Hymenoptera, from Dr.
Krombein’s colleagues around the world. Biographic highlights and a bibliography of Dr.
Krombein’s publications by Beth B. Norden are presented.
Memoir No. 18
Contributions on Diptera, Dedicated to Willis W. Wirth.
Edited by Wayne N. Mathis and William L. Grogan, Jr.
297 pp. 1997 (cost: $25.00).
A memorial volume dedicated to D. Willis W. Wirth, eminent Dipterist and long-time
employee of the Systematic Entomology Laboratory, U.S. Department of Agriculture,
Washington, D.C. A collection of invited papers, mostly on Diptera, prepared by Dr.
Wirth’s colleagues around the world.
Memoir No. 19
Monograph of the Stilt Bugs, or Berytidae (Heteroptera), of the
Western Hemisphere. By Thomas J. Henry. To be published in 1997.
Western Hemisphere Berytidae include 3 subfamilies, 13 genera, and 53 species. Keys
and numerous illustrations are provided for identification, and information on host plants,
feeding habits, and economic importance are provided. An important, comprehensive work
on this family.
Miscellaneous Publication
A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera)
Second Edition, Revised.
By E. Eric Grissell and Michael E. Schauff. To be published in 1997.
This is a revised edition of the 1990 handbook on chalcids by the same authors. The
content is updated with the numerous advances that have appeared since 1990. Pictorial
keys and discussions of the characteristics, biology, distribution, and literature of each
family as well as collecting and mounting techniques and morphology make this a most
useful introduction and overview of the superfamily.
Cynipid Galls of the Eastern United States, by Lewis H. Weld. 124 pp. 1959
Cynipid Galls of the Southwest, by Lewis H. Weld. 35 pp. 1960
PUBLICATIONS FOR SALE BY THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON
MISCELLANEOUS PUBLICATIONS
Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C.
OSL Bayo U0 71 eae ee ee
Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C.
UNS STIRS 0) 9 Sel 8 0? Ie aS aes ES ee Se ee
A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael
PTE SIAN RY 5 wy 2 apa UR eMC Dh a NE EOE SS ae eee ee eee
No.
MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 _____
A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving.
SPS apyes Cie ee eee te ee ee ee
The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman.
SAS iain yd ho Se ee eS Ree ee BR eee ae Se eee
A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952 —-------
A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 __-
The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi
ier EN SOMA La oR C8) ot Bee Te occ SSE ah ob Dn od en
Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp.
ISD Gy pees ak OP Ee I i WO Bl A A a ee ee
The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato-
poponidae); by W. L: Grogan; Jr. and W..W- Wirth. 125 pp. 1979)
The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson.
SA O1UES 9 ofthe Fs) Ry weedeat G/DL ESA eR SN SA ee
. Recent Advances in Dipteran Systematics: Commemorative Volume in Honor of Curtis W.
Sabrosky. Edited by Wayne N. Mathis and F. Christian Thompson. 227 pp. 1982 _._-
. A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370
52 gl AES Ta Aas 2a Co i ee eet eee eee we ee
. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff.
Sg 516 a 2 oN op le NP ie sm oo aeres OOM LENG A Be Seda ea ee
. An Identification Manual for the North American Genera of the Family Braconidae (Hyme-
noptera), by Paul M. Marsh, Scott R. Shaw, and Robert A. Wharton. 98 pp. 1987
. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174
Ss peer tn hae Ee ee a ee
. A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent
HiGwW Grogan, sf:0198 pp) 1999) 2.88. ee ee
. The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 _—
. Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Krombein, edited
by B: B: Norden and A. S. Menke. 216 pp. 1996...»
. Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and
Withimilc Gropan, Jt..297 pp: 1997 2. te
10.00
$15.00
(out of
print)
15.00
15.00
15.00
15.00
11.00
12.00
10.00
11.00
18.00
5.00
18.00
25.00
25.00
25.00
25.00
25.00
Back issues of the Proceedings of the Entomological Society of Washington are available at $60.00 per volume
to non-members and $25.00 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, % Department of Entomology, MRC 168, Smithsonian
Institution, Washington, D.C. 20560.
CONTENTS
(Continued from front cover)
PINTO, JOHN D., RICHARD STOUTHAMER, and GARY R. PLATNER—A new cryptic
species of Trichogramma (Hymenoptera: Trichogrammatidae) from the Mojave Desert of
California as determined by morphological, reproductive and molecular data
THOMPSON, F. CHRISTIAN—Revision of the Eristalis flower flies (Diptera: Syrphidae) of
the Americas south of the United States
TOGASHI, ICHIJI—A new genus and species belonging to the subfamily Sterictiphorinae (Hy-
menoptera: Symphyta: Argidae) from Japan
NOTES
DUFFIELD, RICHARD M: and CHARLES H. NELSON—Note on stoneflies (Plecoptera),
particularly Prostoia besametsa (Ricker) (Nemouridae), in the diet of salmonids from the
headwaters of the middle fork of the South Platte River, Colorado
MENKE, ARNOLD S.—Species homonymy in Belomicrus Costa (Hymenoptera: Sphecidae:
@rabronimae)t open kn woken eae bei oO) se Meeps Oe af. cin dah racatee See tara ade xtae eo tenet eae
SMITH, DAVID R.—Colléections of Stephanidae (Hymenoptera) in the mid-Atlantic states,
including an eastern record for Schlettererius cinctipes (Cresson)
STAINES, C. L.—Cereal leaf beetle (Coleoptera: Chrysomelidae) as a pest of ornamental
grasses
OBITUARY : oh ra ee
SABROSKY, CURTIS W.—George C. Steyskal (1909-1996)
BOOK REVIEW
HORMIGA, GUSTAVO—Riceland Spiders of South and Southeast Asia by A. T. Barrion
and J. A. Litsinger
MISCELLANEOUS
Reports of Officers
Society Meetings
i alee
VOL. 99 JULY 1997 NO. 3
e yi (ISSN 0013-8797)
J
PROCEEDINGS
of the
ENTOMOLOGICAL SOCIETY
of WASHINGTON
SMITHSONGRUBLISHED
JUL $9 99 QUARTERLY
é api ae
CONTENTS
DUSSE, KATHLEEN and L. E. HURD—Food limitation reduces body length in mantid nymphs,
Tenodera sinensis Saussure (Mantodea: Mantidae): implications for fitness ............... ye 490
EASTON, EMMET R. and WING-WAH PUN—Observations on some Hemiptera/Heteroptera
GieViacau Southeast ASIAN, Jo. oi iivdanre few don tyeleiaens shige eae Groin el auc cabioeeerie taconite s ueaainaels stat 574 =
HEADRICK, DAVID H. and RICHARD D. GOEDEN—Gall midge forms galls on fruit fly
palsouipteras© ecidomyidae, Tephritidae) io... cc cene sce cameos oe eieisine ins anit ete sete Slee 487
HEADRICK, DAVID H., RICHARD D. GOEDEN, and JEFFREY A. TEERINK—Taxonomy
of Aciurina trixa Curran (Diptera: Tephritidae) and its life history on Chrysothamnus nau-
seosus (Pallas) Britton in southern California; with notes on A. bigeloviae (Cockerell) ..... 415
JASCHHOF, MATHIAS—Descriptions of a new genus and six new species of Nearctic Lestre-
maitiiaes(Oipteras CECldOmyITdAS)) 2)..4 2552p. pels cts = nate tals sralaiets eestoie Siakelae Sole pate ota sto ole ntores eRefepatolare wae aE ta 523
LARSON, LEN and B. A. FOOTE—Biology of four species of Notiphila Fallén (Diptera:
Ephydridae) associated with the yellow water lily, Nuphar luteum (Nymphaeaceae) ........ 541
LUGO-ORITZ, C. R. and W. P. MCCAFFERTY—A new genus and redescriptions for African
species previously placed in Acentrella (Ephemeroptera: Baetidae) ......................--.--- 429
MARSHALL, S. A.—A first record of Aptilotus Mik (Diptera: Sphaeroceridae) from the Neo-
tropical Region, with the description of three new wingless species of the Aptilotus para-
aoxus eroup trom: hichrelevations: in Costa) Rica 5.22 .))22 je surest icisinncla seen Nets s caters slack 505
MELIKA, GEORGE and WARREN G. ABRAHAMSON—Descriptions of four new species of
cynipid gall wasps of the genus Neuroterus Hartig (Hymenoptera: Cynipidae) with redes-
Y criptions of some known species from the eastern United States ......................0..22285. 560
MILLER, DOUGLASS R. and SRIDHAR POLAVARAPU—A new species of mealybug in the
genus Dysmicoccus (Hemiptera: Coccoidea: Pseudococcidae) of importance in highbush
blueberries (Vaccinium corymbosum, Ericaceae) in the eastern United States ................ 440
MILLER, KELLY B.—Crinodessus amyae, a new Nearctic genus and species of predaceous
diving beetle (Coleoptera: Dytiscidae: Hydroporinae: Bidessini) from Texas, U.S.A. ....... 483
(Continued on back cover)
THE
ENTOMOLOGICAL SOCIETY
OF WASHINGTON
ORGANIZED MARCH 12, 1884
OFFICERS FOR 1997
M. ALMA SOLIs, President MICHAEL G. POGUE, Treasurer
WARREN E. STEINER, JR., President-Elect Davip G. FurTH, Program Chair
HAROLD HARLAN, Recording Secretary JOHN W. BRown, Membership Chair
HOLiis B. WILLIAMS, Corresponding Secretary RALPH P. ECKERLIN, Past President
Gary L. MILLER, Custodian
DAVID R. SmiTH, Editor
Publications Committee
THOMAS J. HENRY WAYNE N. MATHIS
Gary L. MILLER, Book Review Editor
Honorary President
CurTiIs W. SABROSKY
Honorary Members
LOUISE M. RUSSELL ALAN STONE KARL V. KROMBEIN
All correspondence concerning Society business should be mailed to the appropriate officer at the following
address: Entomological Society of Washington, % Department of Entomology, MRC-168, Smithsonian Insti-
tution, Washington, D.C. 20560.
MEETINGS.—Regular meetings of the Society are held in the Natural History Building, Smithsonian Institu-
tion, on the first Thursday of each month from October to June, inclusive, at 8 P.M. Minutes of meetings are
published regularly in the Proceedings.
MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology.
Annual dues for members are $25.00 (U.S. currency).
PROCEEDINGS.—tThe Proceedings of the Entomological Society of Washington (ISSN 0013-8797) are pub-
lished quarterly beginning in January by The Entomological Society of Washington. POSTMASTER: Send
address changes to the Entomological Society of Washington, % Department of Entomology, MRC-168, Smith-
sonian Institution, Washington, D.C. 20560. Members in good standing receive the Proceedings of the Ento-
mological Society of Washington. Nonmember U.S. subscriptions are $60.00 per year and foreign subscriptions
are $70.00 per year, payable (U.S. currency) in advance. Foreign delivery cannot be guaranteed. All remittances
should be made payable to The Entomological Society of Washington.
The Society does not exchange its publications for those of other societies.
PLEASE SEE PP. 615-616 OF THE JULY 1996 ISSUE FOR INFORMATION 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, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash-
ington, D.C. 20560.
Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology,
Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560.
Books for Review: Gary L. Miller, Systematic Entomology Laboratory, ARS, USDA, Building 046, BARC-
West, Beltsville, MD 20705.
Managing Editor and Known Bondholders or other Security Holders: none.
This issue was mailed 11 July 1997
Second Class Postage Paid at Washington, D.C. and additional mailing office.
PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA
This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 409-414
TRIOZA CHENOPODII REUTER: FIRST NORTH AMERICAN RECORD OF
AN OLD WORLD PSYLLID (HOMOPTERA: PSYLLOIDEA: TRIOZIDAE)
A. G. WHEELER, JR. AND E. RICHARD HOEBEKE
(AGW) Department of Entomology, Clemson University, Clemson, SC 29634, U.S.A.;
(ERH) Department of Entomology, Cornell University, Ithaca, NY 14853, U.S.A.
Abstract.—Collection of the psyllid Trioza chenopodii Reuter (Triozidae) at Charlotte-
town, Prince Edward Island, Canada, in July 1995, represents the first record of this
common Palearctic species in North America. Adults were collected on Atriplex littoralis
L. (Chenopodiaceae). A summary of this psyllid’s Old World distribution, the biological
literature, and a taxonomic diagnosis and description are provided.
Key Words:
Homoptera, Psylloidea, Trioza chenopodii, Triozidae, detection, adventive
insects, Chenopodiaceae, Atriplex
Our work on adventive insects in eastern
North America has emphasized their detec-
tion in areas especially vulnerable to inva-
sion and establishment by Old World spe-
cies, documentation of their spread, and ob-
servations on their seasonality and habits.
Since 1993, we have collected in the Mar-
itime Provinces of Canada, which histori-
cally have provided numerous records of
immigrant insects (e.g., Brown 1940, Mor-
ris 1983, Hoebeke and Wheeler 1996).
Nova Scotia, in particular, continues to
yield species new to the Nearctic Region
(see Hoebeke and Wheeler 1996). Herein
we report the first North American collec-
tion of the psyllid Trioza chenopodii Reuter
(Triozidae); summarize the Old World lit-
erature on its distribution, life history, and
habits; and provide a diagnosis and descrip-
tion to facilitate its recognition among other
psyllids in the Nearctic fauna.
TAXONOMIC HISTORY
Trioza chenopodii is a polymorphic spe-
cies that occurs in both a longwinged sum-
mer form (aestivalis) and a shortwinged au-
tumn and winter form (autumnalis). These
forms, long considered separate species,
were established through laboratory rearing
to represent dimorphic forms of a single
species (Lauterer 1982). Originally de-
scribed in Trioza Forster (Reuter 1876),
chenopodii was placed in the subgenus Het-
erotrioza (Dobreanu and Manolache 1962),
which was raised to generic level by Kli-
maszewski (1968). In that taxonomic
scheme, chenopodii belongs to the nomi-
nate subgenus Heterotrioza (Klimaszewski
1973). Here, however, we follow Hodkin-
son and White (1979) and Ossiannilsson
(1992) in retaining this species in Trioza.
DIAGNOSIS AND DESCRIPTION
Trioza chenopodii Reuter
(synonymy: after Lauterer, 1982:148;
Ossiannilsson, 1992:324)
Trioza chenopodii Reuter, 1876:76 (= form
autumnalis).
Trioza obliqua Thomson, 1877:825 (=
form aestivalis).
Trioza dalei Scott, 1877:31 (= f. autum-
nalis).
Trioza atriplicis Lichtenstein, 1879:cxv (=
f. aestivalis and ? partly f. autumnalis).
410
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-2.
= 0.5 mm and for Fig. 2 = 1.0 mm. (From Ossiannilsson, 1992: 323).
Trioza galii; Jacobsen, 1919:359, nec Fors-
ter, 1848:87 (form indeterminate).
Trioza horvathi; Jacobsen, 1919:360, nec
Low, 1881:263 (= f. aestivalis).
Trioza (Heterotrioza) chenopodii; Klimasz-
ewski, 1973:241.
Heterotrioza chenopodii; Lauterer,
148.
1982:
Trioza, a diverse and ubiquitous genus of
worldwide distribution (Hollis 1984), in-
cludes 45 Nearctic species (Hodkinson
1988). Trioza chenopodii can be readily
distinguished from all other Nearctic psyl-
lids by its preference for the Chenopodia-
ceae (only two Nearctic species of Cras-
pedolepta are known to feed on chenopods;
Hodkinson, 1988:1227) and by its highly
characteristic male genitalia (Figs. 3-5).
This newly detected chenopod specialist
can be recognized as a member of the genus
Trioza (Triozidae) by the following com-
bination of characters: antenna cylindrical,
without long, dark setae; head in dorsal
view with vertex short, at most 0.8 times as
long as broad, and generally much shorter;
genae with conspicuous processes (genal
cones) (Fig. 1); forewing with veins Cul
and M not having a common stem, each
arising separately from common origin at
vein R (Fig. 2); and forewing without
brown color pattern, occasionally uniformly
yellow. The following description is taken
from Hodkinson and White (1979:75) and
Ossiannilsson (1992:324).
Adult.—Extremely variable in color;
Trioza chenopodii, male (f. aestivalis). 1, Head, frontal aspect. 2, Forewing. Scale line for Fig. 1
young specimens generally yellow or yel-
low-green with brown markings, abdomen
often green; in older specimens dorsum of-
ten uniformly dark brown. Antenna varying
in pigmentation, but generally segments
II-V pale, remaining segments brown or
black. Seasonally dimorphic: autumn-win-
ter form (autumnalis) with short forewings
and summer form (aestivalis) with fore-
wings normal in length. Metatibia with 3
(arranged 2 + 1) thick, black apical spines.
Forewing with surface spinules largely con-
fined to basal half of wing. Shape and sur-
face structure of head as in Fig. 1. Male
genitalia (Figs. 3-5): proctiger in lateral as-
pect with well-developed posterior lobe
(Fig. 3); paramere in lateral aspect tapering
to acute apex (Fig. 4); apical portion of ae-
deagus with highly characteristic subapical
process (Fig. 5). Female terminalia (Figs.
6—7): proctiger in lateral and dorsal aspects
long (Figs. 6—7); circum-anal ring as in Fig.
7. Overall length of male (European litera-
ture) 2.10—2.71 mm (form aestivalis), of fe-
male (European literature) 2.48—3.05 mm
(form aestivalis). Principal hosts: Cheno-
podium, Atriplex, and Halimione (European
literature).
For a description of the fifth-instar
nymph of both forms (aestivalis and autum-
nalis), the reader is referred to Ossiannils-
son (1992:324—325).
GEOGRAPHIC DISTRIBUTION
Trioza chenopodii is widespread in the
Palearctic Region, occurring in Great Brit-
VOLUME 99, NUMBER 3 411
Figs. 3-5. Trioza chenopodii, male (f. aestivalis). 3, Terminalia, lateral aspect. 4, Left paramere, lateral
aspect. 5, Apical portion of aedeagus (showing subapical process), lateral aspect. Scale lines = 0.1 mm. (From
Ossiannilsson, 1992: 324).
ain, most of continental Europe, Canary Is- ewski 1973, Lauterer 1982, Burckhardt
lands, Cyprus, the Caucasus and other por- 1986, Hodkinson and White 1979, Ossian-
tions of the former USSR, North India, Pa- nilsson 1992),
kistan, Iran, Israel, and Egypt (Klimasz- On July 26, 1995, adults of the long-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 6—7.
anal ring), dorsal aspect. Scale lines = 0.1 mm. (From Ossiannilsson, 1992: 325).
winged form (aestivalis) were found on
Atriplex littoralis L. (= A. patula var. lit-
toralis (L.) Gray) growing along the shore
at Charlottetown, Prince Edward Island.
The 36 adults collected are deposited in the
Cornell University Insect Collection, Itha-
ca, NY (2192, 9d), and the National Mu-
seum of Natural History, Beltsville, MD
GPz,.3:5.)
BIONOMICS
This psyllid is common in ruderal habi-
tats, often between 160 and 300 meters
above sea level, where it develops on var-
ious Chenopodiaceae. Species of Cheno-
podium and Atriplex are the principal hosts
(Lauterer 1982, Burckhardt 1986), although
it occurs on other chenopods, including
Halimione portulacoides (L.) Aell. (Hod-
kinson and White 1979). In the laboratory
it has been reared on sugar beet (Beta vul-
garis L.) and spinach (Spinacea oleracea
L.), although considerable mortality oc-
curred on the latter plant. Early-instar
nymphs are gall formers, feeding within
leaf folds on their hosts, but fourth and fifth
Trioza chenopodii, female (f. autumnalis). 6, Terminalia, lateral aspect. 7, Proctiger (and circum-
instars are free feeders on leaves, stems,
petioles, and inflorescences (Lauterer
1982). Trioza chenopodii is multivoltine
over much of its range but apparently pro-
duces only two generations in Great Britain
and Scandinavia (Hodkinson and White
1979, Lauterer 1982, Ossiannilsson 1992).
However, in the plains and foothills of
northern Punjab (Rawalpindi area), Paki-
stan, at least five generations occur on Che-
nopodium spp. between January and Octo-
ber. There, 7. chenopodii is one of three
insects and a fungus that showed great po-
tential as biological control agents of weedy
Chenopodium in cultivated cereal and veg-
etable fields (Baloch and Ghaffar 1984).
In the former Czechoslovakia, adults of
the shortwinged form overwinter in leaf lit-
ter and grasses near the host plants and
move onto chenopods in late April or early
May. Eggs are laid singly, usually on upper
or lower leaf surfaces. Adults of the first
summer generation appear from late May to
early June. Adults of the first and second
summer generations overlap during August.
The season’s final generation, comprised of
VOLUME 99, NUMBER 3
shortwinged adults induced by decreasing
daylength, may not appear until early No-
vember. Thus, Czechoslovakian populations
typically consist of two (rarely three) sum-
mer (aestivalis) generations and one autum-
nalis generation (Lauterer 1982). For addi-
tional biological information, including the
effects of photoperiod on diapause, the
reader is referred to Lauterer’s study.
Atriplex littoralis, on which T. chenopo-
dii was collected in Prince Edward Island,
is also recorded as a host of this psyllid in
Europe (Hodkinson and White 1979, Lau-
terer 1982). This Holarctic chenopod, wide-
spread in North Temperate regions, is re-
garded as an obligate halophyte (Tascher-
eau 1985). In northeastern North America,
diploid coastal populations of A. littoralis
are thought to be mainly introduced, where-
as hexaploid plants occurring in inland sa-
line habitats (and occasionally along the
coast) are probably native (Gleason and
Cronquist 1991).
MODE OF INTRODUCTION
We consider 7. chenopodii to have been
introduced accidentally into North America,
perhaps with ship’s ballast, nursery stock,
or packing material used in shipping plants.
More than 20 other psyllids are believed to
be adventive in North America (Downer et
al. 1988; Hodkinson 1988; Gill 1987, 1991;
Pozzi 1988). Trioza chenopodii, which ap-
pears to be overlooked frequently in Europe
(Ossianilsson 1992), might be more wide-
spread in coastal populations of A. littoralis
in northeastern North America.
ACKNOWLEDGMENTS
We express our appreciation to G. L.
Miller (USDA-ARS, Systematic Entomol-
ogy Laboratory, Beltsville, MD) for veri-
fying our identification of Trioza chenopo-
dii, for reviewing the manuscript, and for
providing useful comments for improve-
ment, and to P. H. Adler and J. C. Morse
(Clemson University, Clemson, SC) for re-
viewing the manuscript. We also acknowl-
edge the publisher, E. J. Brill (Leiden, The
413
Netherlands), for permission to reproduce
Figs. 1-7 from Ossiannilsson’s 1992 paper.
This is Technical Contribution No. 4236 of
the South Carolina Agricultural Experiment
Station, Clemson University.
LITERATURE CITED
Baloch, G. M. and A. Ghaffar. 1984. Natural enemies
of Chenopodium spp. in Pakistan with notes on
Trioza chenopodii |[Hem.:Psyllidae], a promising
biocontrol agent. Entomophaga 29: 409-414.
Brown, W. J. 1940. Notes on the American distribu-
tion of some species of Coleoptera common to the
European and North American continents. Cana-
dian Entomologist 72: 65-78.
Burckhardt, D. 1986. Redescription of Trioza sahl-
bergi Sulc with comments on the genus Hetero-
trioza (Hem.-Hom., Psyllidae). Entomol. Mon.
Mag. 122: 121-126.
Dobreanu, E. and C. Manolache. 1962. Homoptera
Psylloidea. [In Rumanian]. Fauna République Po-
pulaire Roumaine, Bucharest. Insecta 8(3): 1-376.
Downer, J. A., P. Svihra, R. H. Molinar, J. B. Fraser,
and C. S. Koehler. 1988. New psyllid pest of Cal-
ifornia pepper tree. California Agriculture 42(2):
30-31.
Forster, A. 1848. Ubersicht der Gattungen und Arten
in der Familie Psylloden (mit 2 Nachtragen). Ver-
handlungen des Naturhistorischen Vereins der
Preussichen Rheinlande 5: 65-98.
Gill, R. J. 1987. “Acizzia acaciaebaileyanae.”’ Detec-
tion Advisory PD13-87, California Department of
Food Agriculture, Sacramento, CA, 8 June, 1987.
Gill, R. J. 1991. “Blue gum psyllid.”” Detection Ad-
visory PDO3-91. California Department of Food
Agriculture, Sacramento, CA, 25 February, 1991.
Gleason, H. A. and A. Cronquist. 1991. Manual of
Vascular Plants of Northeastern United States and
Adjacent Canada, Second Edition. The New York
Botanical Garden, Bronx, NY. 910 pp.
Hodkinson, I. D. 1988. The Nearctic Psylloidea (In-
secta: Homoptera): an annotated check list. Jour-
nal of Natural History 22: 1179-1243.
Hodkinson, I. D. and I. M. White. 1979. Homoptera
Psylloidea. Handbooks for the Identification of
British Insects, Vol. If, Part 5(a): 198.
Hoebeke, E. R. and A. G. Wheeler, Jr. 1996. Meli-
gethes viridescens (F.) (Coleoptera: Nitidulidae) in
Maine, Nova Scotia, and Prince Edward Island:
diagnosis, distribution, and bionomics of a Pale-
arctic species new to North America. Proceedings
of the Entomological Society of Washington 98:
221-227.
Hollis, D. 1984. Afrotropical jumping plant lice of the
family Triozidae (Homoptera: Psylloidea). Bulle-
tin of the British Museum (Natural History) (En-
tomology) 49: 1-102.
414
Jacobsen, O. 1919. Fortegnelse over de hidtil kendte
danske Psyllider. Entomologiske Meddelelser 12:
355-360.
Klimaszewski, S. M. 1968. Relations de parenté des
espéces du genre Trioza Forst. (Homoptera, Psyl-
lodea) de l’Europe centrale 4 la lumiére des exa-
mens faits par les méthode de taxonomie numé-
rique. [In Polish, French summary]. Annales
Universitatis Mariae Curie-Sklodowska, Sect. C,
22: 1-20.
Klimaszewski, S. M. 1973. The jumping plant lice or
psyllids (Homoptera, Psyllodea) of the Palearctic:
an annotated check-list. Annales Zoologici, War-
szawa 30: 155-286.
Lauterer, P 1982. New data on the occurrence, bio-
nomics and taxonomy of some Czechoslovakian
Psylloidea (Homoptera). Acta Musei Moraviae 67:
133-162.
Lichtenstein, M. J. 1879. Quelques observations en-
tomologiques—Trioza atriplicis (sp. n.). Bulletin
des Séances de la Société Entomologique de
France, Ser. 5, t. 9: cxiv—cxv.
Low, E 1881. Beschreibung von zehn neuen Psyllo-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
den-Arten. Verhandlungen der Zoologische-Bo-
tanische Gesellschaft in Wien 31: 255-268.
Morris, R. E 1983. Introduced terrestrial insects, pp.
551-591. In South, G. R., ed., Biogeography and
Ecology of the Island of Newfoundland. W. Junk,
The Hague, Netherlands.
Ossiannilsson, E 1992. The Psylloidea (Homoptera)
of Fennoscandia and Denmark. Fauna Entomolo-
gica Scandinavica, Vol. 26. E. J. Brill, Leiden,
Netherlands. 347 pp.
Pozzi, J. 1988. ‘‘Psyllid.”” Detection Advisory PD49-
88. California Department of Food Agriculture,
Sacramento, CA, 6 September, 1988.
Reuter, O. M. 1876. Catalogus Psyllodearum in Fen-
nia hactenus lectarum. Meddelanden af Societas
pro Fauna et Flora Fennica 1: 69-77.
Scott, J. 1877. Description of a new species of Trioza.
Entomologist’s Monthly Magazine 14: 31—32.
Taschereau, P. M. 1985. Taxonomy of Atriplex species
indigenous to the British Isles. Watsonia 15: 183—
209.
Thomson, C. G. 1877. Ofversigt af Skandinaviens
Chermes-arter. Opuscula Entomologica 8: 820—
841.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 415-428
TAXONOMY OF ACIURINA TRIXA CURRAN (DIPTERA: TEPHRITIDAE)
AND ITS LIFE HISTORY ON CHRYSOTHAMNUS NAUSEOSUS (PALLAS)
BRITTON IN SOUTHERN CALIFORNIA; WITH NOTES ON
A. BIGELOVIAE (COCKERELL)
Davip H. HEADRICK, RICHARD D. GOEDEN, AND JEFFREY A. TEERINK
Department of Entomology, University of California, Riverside, CA 92521, U.S.A.
Abstract.—Aciurina trixa Curran is reinstated as a valid taxon and distinguished from
A. bigeloviae (Cockerell) with which it had been synonymized. The immature stages of
A. trixa also are described in detail. This tephritid is univoltine and monophagous on
Chrysothamnus nauseosus (Pallas) Britton. Adults emerge in early spring and lay eggs
singly in axillary buds on the current season’s branches. First instars eclose in ca. 1 week
and tunnel basipetally along a vascular trace into a branch. Gall growth begins during the
first larval stadium and continues through the spring and summer. Gall growth was com-
plete by late fall when most branch tunnels and galls contain second instars. Second instars
continue to feed in the branch tunnels and begin excavating the gall lumen. Larval growth
proceeds through the fall, and by December most galls contain third instars. Third instars
confined most of their feeding to the gall, typically filling the branch tunnel with packed
frass. Puparia were first observed in mid-February. Pupariation took place within the gall
lumen and lasted ca. 2 weeks. Adult females were proovigenic and mated shortly after
emergence and repeatedly.
Key Words:
parasitoids
Aciurina trixa Curran is reinstated as a
valid species, the immature stages are de-
scribed for the first time, and a redescription
of the adult is provided. The biology, ovi-
position behavior, distribution, and natural
enemies of A. trixa are also described for
populations occurring in southern Califor-
nia.
MATERIALS AND METHODS
Our field studies of the biology of A.
trixa were conducted between 1989 and
1994 in southern California. Our primary
study and collection locations were Los An-
geles County, Pearblossom at ca. 830 m;
San Bernardino Co., Cajon Junction at ca.
1000 m, and Pioneertown at ca. 1200 m.
Insecta, Aciurina taxonomy, immature stages, galls, oviposition behavior,
Field studies of adult behaviors were con-
ducted at the Cajon Junction site during the
spring of 1990-1992 on C. nauseosus.
Field data were supplemented by laboratory
dissections, measurements, and photogra-
phy of infested host-plant material, and sub-
sequent rearing of adults and their parasit-
oids.
Twenty-three ova were obtained for
study by dissection of gravid, field-collect-
ed females. Five first, 20 second, and 18
third instar larvae and two puparia were
dissected from infested C. nauseosus and
all immature stages were examined using
scanning electron microscopy (SEM). All
specimens for SEM were fixed in 70%
EtOH, rehydrated to distilled water in a de-
416
creasing series of acidulated EtOH, post-
fixed in 2% aqueous osmium tetroxide for
24 h, dehydrated to absolute EtOH in an
increasing series of EtOH dilutions, critical-
point dried, mounted on stubs, sputter-coat-
ed with a gold-palladium alloy and exam-
ined on a JEOL JSM C35 scanning electron
microscope in the Department of Nematol-
ogy, University of California, Riverside.
Micrographs were prepared on Polaroid
P/N 55 film at 15 kV accelerating voltage.
Tephritid names follow Foote et al.
(1993), except where amended below; lar-
val terminology follows Headrick and Goe-
den (1991) and Goeden and Teerink (1996a,
b, c); terms for adult behaviors follows
Headrick and Goeden (1994); and plant
names follow Munz (1968, 1974) and
Munz and Keck (1959). Abbreviations used
as follows: AMNH-—American Museum of
Natural History; CAS—California Academy
of Sciences; CDFA-—California Department
of Food and Agriculture; UIM—University
of Idaho, Moscow; UCR—University of Cal-
ifornia, Riverside; WSUP-—Washington
State University, Pullman. Voucher speci-
mens of adult A. trixa and its parasitoids
reside in the research collection of RDG.
Preserved immature stages are held in the
research collection of JAT. Means + stan-
dard errors are provided throughout.
RESULTS AND DISCUSSION
TAXONOMY
The taxonomy of Aciurina species oc-
curring in the western United States has had
a tortuous history. Unclear or unknown
host-plant associations, variable gall mor-
phology, incorrect insect gall-former attri-
butions, and the recently discovered region-
al sexual dimorphisms (Goeden and Teerink
1996b, c) have all contributed to the diffi-
culty of elucidating the species comprising
this genus. Based on our extensive collec-
tions of reared material and examination of
museum specimens, many taxonomic issues
have recently been clarified (Goeden and
Teerink 1996a, b, c).
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Although, A. trixa has been mentioned in
the literature subsequent to its synonymi-
zation with Aciurina bigeloviae (Cockerell),
no formal resurrection of this species has
been published. Most recently, notes on the
biology, gall formation, and sexual behav-
ior of A. trixa were reported from northern
Arizona by Fernandez and Price (1994),
who cited the work of Dodson and George
(1986) for identification of the species that
they studied. Foote et al. (1993) remarked
that the names synonymized by Steyskal
(1984) and Norrbom (1989) were available.
Adult.—Curran (1932) erected the genus
Aciurina and designated the type species as
A. trixa. Steyskal (1984) revised the genus
and synonymized A. semilucida (Bates)
(1935) and A. trixa with A. bigeloviae
(Cockerell). Recently, Goeden and Teerink
(1996c) resurrected A. semilucida based on
the discovery of sexually dimorphic wing
patterns among disparate populations there-
tofore unrecognized. This discovery was
based primarily on examination of flies in-
dividually reared from known galls which
reconfirmed the separate species status of A.
semilucida. Similarly, Dodson and George
(1986) provided biological, behavioral, and
genetic evidence that A. trixa was distinct
from A. bigeloviae based on flies individu-
ally reared from known galls from through-
out the southwestern U.S., excluding south-
ern California, but they did not formally
resurrect the former species. Our examina-
tion of material, reared or otherwise col-
lected, in southern California and museum
specimens from all states west of the Rock-
ies, except Oregon, provides more pieces of
the puzzle concerning the taxonomic status
of A. trixa.
The following description supplements
the original description of Curran (1932),
with characters since recognized as impor-
tant in separating Aciurina species (Steys-
kal 1984, Foote et al. 1993).
Aciurina trixa Curran
(Fig. 61)
Female holotype. Head.—White to gold-
en; antenna golden, arista lighter basally,
VOLUME 99, NUMBER 3
darker apically; bristles as in original de-
scription; all uniformly concolorous with
head basally, but dark apically.
Thorax.—Dark gray, with uniform cov-
ering of gray pollinosity, setulae white.
Bristles as given in original description,
their location as given in Foote et al.
(1993). Scutellum dark brown with poste-
rior apex lighter. Coxae and legs golden;
fore-coxa with patch of very fine setae an-
teriorly; forefemur thickened dorsally, with
2-3 rows of short stout setae, a ventrolat-
eral row of longer setae extending entire
length of femora, and at most a single in-
complete row of dark setae ventrally (see
male description). Wing mostly dark brown
with marginal hyaline incisions and internal
markings as illustrated by Curran (1932
Fig. 5) (see below for variations). Pterostig-
ma with a proximal hyaline incision; hya-
line spot in cell br round; cell cua, with 1
large and 2 small marginal hyaline inci-
sions, larger one includes anal lobe; cell m
with 2 marginal hyaline incisions, distal in-
cision extends into cell r,,;, often touching
vein R,,;.
Abdomen.—Shiny, reddish brown,
sparsely covered with very fine yellow bris-
tles, tergite 6 darker, at least in patches, than
preceding segments; syntergosternite 7 fus-
cous to black; aculeus (visible in type) lan-
ceolate without distinct markings or inden-
tations.
Male. Head.—White to golden, vertex
darker, ocellar triangle dark; bristles stout,
golden; antenna golden, arista golden ba-
sally, dark apically.
Thorax.—Dark, fine gray pollinose, setu-
lae short white, dorsocentral seta nearer to
postsutural supralar seta than in female;
legs golden; fore coxa with anterior patch
of fine setae; forefemur thickened, much
larger than in remaining legs, larger than
female forefemur relative to sizes in re-
maining legs; rows of stout setae dorsally,
ventrolateral row of larger setae extending
entire length of femur; ventrally 2—3 rows
of short dark setae (possibly aid in securing
hold on female during copulation [Dodson
417
1987]). Wing with marginal hyaline inci-
sions and internal markings similar to that
of female (see variations below).
Abdomen.—Brown with darker mark-
ings anteriorly, dark around lateral margins,
tergite 1 almost completely dark, tergite 5
dark laterally; post-abdomen dark, surstyli
golden, prensisetae dark.
Variation.—In males, the head color var-
ies from golden to brown. The setulae of
the thorax in both sexes are white or or-
ange. Variation in setulae color among pop-
ulations of flies was not observed; most
specimens, including the holotype of A.
trixa, have the white form. In California,
the specimens reared from small galls all
have orange setulae. The thorax color in
both sexes varies from light brown to nearly
black.
The wing markings varied among the
specimens examined. The hyaline or sub-
hyaline incision on the proximal side of the
pterostigma can be quite faint among the
specimens reared from California. This was
also observed among specimens collected
(swept or reared) from Idaho. The individ-
uals reared from small galls in southern
California were darker than those reared
from the larger galls and the infuscation of
their wings was heavy. However, even in
these flies, a faint mark on the proximal
side of the pterostigma was observed, es-
pecially on or near the costal vein. The hy-
aline spot in cell br varies from circular to
oblong in A. trixa. Only very rarely will
specimens (both sexes) have a small infus-
cation on the wing margin within the prox-
imal hyaline mark in cell m, but it is not
elongate and never complete (i.e. bridging
the infuscation, thereby forming 2 hyaline
incisions from the proximal hyaline mark).
In cell c the infuscation can be either nar-
row or wide with the widest marking being
nearly equal in width to the distance be-
tween vein S. and R,. In approximately 4%
of the female specimens examined, the
wings have three hyaline marks in cell r,
like A. trilitura Blanc and Foote, with only
the distal mark either present or absent. Cell
418
cua, typically has 2 small hyaline markings
distally and a larger one proximally. The
distal hyaline mark in cell m is sometimes
constricted at vein M; in a few specimens
the constriction is complete and separates
the mark in cell m from the mark in cell
r,,5- The hyaline mark in cell r,,; can also
be constricted and thus bilobed. The mark
touches vein R,,; in about half of the spec-
imens examined. In 3 specimens examined,
the hyaline mark in cell r,,; is completely
constricted forming 2 distinct marks.
Material examined.—Aciurina trixa: Fe-
male holotype, UTAH: Stansbury L.,
13.vi.13 (AMNH). CALIFORNIA: Inyo
Co., Grapevine Canyon, 2 males | female,
20.v.1988, (R. D. Goeden) (UCR); Lone
Pine, 1 male, no gall, 1.vi.1937, (E. P. Van
Duzee) (CAS). Kern Co., Cane Brake Val-
ley, W. Side Walker Pass, 3,000 ft., 1 fe-
male and | male with gall, 1 female without
gall, Chrysothamnus nauseosus, 9.xi.70, (J.
T. Howell) (CAS); Spanish Needle Creek,
3 females with galls, 6 males with galls,
11.iv.1989, (R. D. Goeden) (UCR); Walker
Pass, 1 male with gall, 11.1v.1989, (R. D.
Goeden) (UCR). Los Angeles Co., Pear-
blossom, 7 males with galls, 5 females with
galls, 18.11.1987, (R. D. Goeden) (UCR).
Maricopa, Co., 9 females, 2 males,
31.11.1931, (E. PB VanDuzee) (CAS). San
Bernardino, Co., Barstow, 1 female, ‘Ex.
Artemisia’, 16.iv.1938, (T. G. H. Aitken)
(CAS); Forest Falls, 7 females with galls, 2
males with galls, 3.v.1988, (R. D. Goeden)
(UCR). San Luis Obispo Co., Maricopa
Grade, 4 males with galls, 3 females with
galls, 21.111.1931, (E. PR VanDuzee) (CAS).
Santa Barbara Co., Ventucopa, 2 females
with: galls, 25.4111.1959, (CR. P.. Allen)
(CDFA). COLORADO: Grand Junction, 1
female! io 20.vi.01e © QWSUP)) >Salidat
28.vi.1933, 3 females 3 males, ‘‘ex Chrys-
othamnus’’, (Cockerell) (WSUP). IDAHO:
All with galls: Bear Lake Co., Dingle, 1
female, sh7iv1. 1975; GW: Teo Wanebere)
(UIM). Boise Co., Horseshoe Bend, 3 fe-
males, 21.v.1975, (J. T. Wangberg) (UIM).
Box Elder Co., Strevell, 1 male, “C. nau-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
seosus’’, 19.i11.1974, (J. T. Wangberg)
(UIM). Butte Co., Howe, 2 males, ““C. nau-
seosus”, 2700-67 (203) Allen): (CUI.
Franklin Co., Preston, 1 male, 18.i11.1974,
(J. T. Wangberg) (UIM). Idaho Co., Lucille,
1 male, 2 females, 12.iv.1975; 2 males,
24.111.1974; 2 females, 28.iv.1974, (Wang-
berg) (UIM); Riggins, 1 female, 29.iv.1974,
(J. T. Wangberg) (UIM); Slate Creek, 1
male, 20.v.1975, (J. T. Wangberg) (UIM).
Lemhi Co., Leadore, 1 female, 2 males,
21.vi.1975, (J. T. Wangberg) (UIM). Oneida
Co., Stone; 1 female, “Co nauscosus™
19.111.1974, (J. T. Wangberg) (UIM). Owy-
hee Co, Walters Ferry, “C. nauseosus’’, 5
females, 3 males, 31.111.1967, (E. J. Allen)
(UIM). Bruneau, 1 female, 26.vi.1974, (J.
T. Wangberg) (UIM). Twin Falls Co., Twin
Falls, 1 female, 29.111.1967, (E. J. Allen)
(UIM). NEVADA: Washoe Co., Hills just
N. of Sparks, 1 male, 1 female, 1 gall, (J.
T. Howell) (CAS). NEW MEXICO: Santa
Fe, 2 males, 21.vi.1926, (EZ. C. VanDyke)
(CAS). UTAH: Provo, 1 female, 1.v.1963,
(WSUP). Female Paratype: Collinston,
21.vii.1927, ‘“‘Det. Curran”? (G. E Knowl-
ton) (AMNH). WASHINGTON: Okanogah
Co., 1 female, no gall, 19.v.1968, (WSUP).
Whitman Co., Steptoe Cyn., 1 female, no
gall, 17.v.1973, (WSUP).
Aciurina bigeloviae. ARIZONA: Flag-
staff, 1 male, no gall, 28.vi.1971, (E. Klee)
(CAS). NEVADA: Reno, 1 female with
gall, 9.v.1891, (E Hilman) (CAS).
Diagnosis—Aciurina trixa is variable in
its adult morphology and the morphology
of the galls it induces on its host plant,
Chrysothamnus nauseosus. Aciurina trixa
can be distinguished from its closest rela-
tive, A. bigeloviae, based primarily on gall
type, and for the majority of specimens by
the presence of a hyaline mark on the prox-
imal side of the pterostigma and by the ab-
sence of an elongate or dividing mark with-
in the proximal hyaline spot in cell m, [see
also Fig. 1 in Dodson and George (1986)].
This distinction makes A. bigeloviae a rare
species known only from large cottony
galls and restricts its range to exclude Cal-
VOLUME 99, NUMBER 3
ifornia. However, one record indicates its
range may extend into northeastern Cali-
fornia along the edge of the Great Basin
(Foote and Blanc 1963).
Aciurina trixa differs from A. semilucida
by having multiple hyaline spots in cell r,,,
(cf. Goeden and Teerink [1996c], Fig. 1A—
E) or from females of A. semilucida occur-
ring in California by not having the distal
hyaline spot in cell m extend beyond vein
R,,; into cell r,,, (cf. Goeden and Teerink
[1996c], Fig. 1F). Aciurina semilucida is
known to form galls on Chrysothamnus vis-
cidiflorus (Hooker) Nuttall and C. nauseo-
sus (Foote et al. 1993). Aciurina semilucida
galls are readily distinguishable from galls
of A. bigeloviae and A. trixa by being light
green, pyriform with an attenuated apex,
covered with a uniform, light-colored pu-
bescence, and bearing several basal leaves
(Goeden and Teerink 1996c).
Both A.
known only from C. nauseosus; however,
the galls they induce are readily distin-
guishable and is the best character for spe-
cies recognition. Aciurina bigeloviae is
known to form only large, cottony galls
[see also Dodson and George (1986), Fig-
ure 2B] that sometimes bear small leaves
studding the gall body, rather than a rosette
of basal leaves as in A. semilucida (Goeden
and Teerink 1996c). The tomentum of A.
bigeloviae galls is thick and continuous
over the entire gall, appearing like a wad of
cotton. Aciurina bigeloviae galls have been
collected from several sites in Idaho, Col-
orado, Arizona, New Mexico and Utah;
however, most specimens in collections do
not have the galls from which they were
reared. The galls of A. maculata (Cole) also
occur on C. nauseosus and are cottony;
however, they are much smaller than A. bi-
geloviae galls and the adults are readily dis-
tinguishable (Foote et al. 1993).
No cottony tephritid galls have ever been
found on C. nauseosus in southern Califor-
nia (RDG, unpublished data). The galls of
A. trixa vary in external morphology
throughout its range. At present we can dis-
trixa and A. bigeloviae are
419
tinguish two types of galls in southern Cal-
ifornia. There are reportedly three gall
forms in Idaho (Wangberg 1981). Galls in
southern California have a waxy or resinous
outer layer. The most common whitish
(pearly) gall found in California is shown
in Fig. 6B. The smaller, brownish gall has
a more restricted distribution, occurring on
the north slope of the San Bernardino
Mountains bordering the Mojave Desert
(shown dissected in Fig. 6G, H).
The key to species of Aciurina as given
in Foote et al. 1993 is modified to include
the resurrection of A. trixa (figures cited are
from Foote et al. 1993, except as noted).
10. Pterostigma along costa no more than 1.5
times as long as its greatest width (Fig. 121,
c); vein dm-cu nearly straight (Fig. 121, e),
the lower apical angle of cell dm about 65°
(Fig. 121, f); wing predominantly hyaline . .
ERE ORET STG cs Cite ae eL eeee e notata (Coquillett)
Pterostigma along costa at least 2.0 times as
long as its greatest width (Fig. 124, a); vein
dm-cu usually bowed apicad (Fig. 124, b), the
lower apical angle of cell dm seldom less than
90° (Fig. 124, c); wing patterned
11. Galls without tomentum; pterostigma of at
least one wing with a proximal hyaline or
subhyaline incision (can be very faint and
only on and slightly extending from costa)
(Fig. 124; Dodson and George, Fig. 1c); prox-
imal marginal hyaline incision in cell m usu-
ally lacking median, dark mark trixa Curran
Galls with cottony tomentum, pterostigma
and costa dark from Sc to R, in most speci-
mens (Fig. 122, b); proximal marginal hyaline
incision in cell m usually with a median, dark,
often elongate mark (Fig. 122), which some-
times divides the incision (Steyskal 1984, Fig.
13) (Dodson and George 1986; Fig. la, b, d)
bigeloviae (Cockerell)
Immature stages. Egg.—White, elongate-
cylindrical with narrow, parallel-sided,
short pedicel and bluntly rounded, basal end
(Fig. 1A). Pedicel with ring of multipored
aeropyles and micropyle at apex (Fig. 1B).
Chorion with slightly elevated striations on
pedicel.
First instar—Translucent white, globose,
bluntly rounded anteriorly and posteriorly,
three to four rows of acanthae circumscribe
420
1SKVY X1i66
Fics ih
pedicel.
Egg of Aciurina trixa. (A) Habitus; (B)
the body at intersegmental lines, lateral
midline with up to four rows of acanthae
(Fig. 2A). Gnathocephalon conical, rugose
pads absent (Fig. 2B). Mouth hook with
two teeth (Fig. 2B-1), cushioned by an in-
tegumental petal (Fig. 2B-3), median oral
lobe present (Fig. 2B-2). Anterior sensory
lobe with four sensory organs (Fig. 2B-4),
dorsal sensory organ comprised of single
papilla (Fig. 2B-5). Stomal sense organ la-
terad of mouth lumen bearing sensory pa-
pillae (Fig. 2B-6). Posterior spiracular
plates contiguous, rounded, and protuding;
each plate bears two rimae (Fig. 2C-1) and
typically four, spine-like, interspiracular
processes (Fig. 2C-2). Compound sensilla
ventrad of spiracle each comprised of two
stelex sensilla (Fig. 2C-3).
Second instar.—White, oblong-elliptical,
bluntly rounded anteriorly and posteriorly,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
16KV
renee g
a524.> 14.00 ° C
Fig. 2. First instar of A. trixa. (A) Habitus, anterior
to left; (B) gnathocephalon, lateral view, 1—mouth-
hooks, 2—median oral lobe, 3—integumental petal,
4—anterior sensory lobe, 5—dorsal sensory organ,
6—stomal sense organ; (C) caudal segment, 1—rima,
2—interspiracular process, 3—compound sensilla.
kegs a
15SkKU"-x1198 ~
VOLUME 99, NUMBER 3
15KV
* Se
15KU ¥2206 @o12 16.80 15kU, 41500 oP Saii-oeot
Fig. 3. Second instar of A. trixa. (A) Habitus, anterior to left; (B) gnathocephalon, lateral view, 1—mouth-
hook, 2—median oral lobe, 3—dorsal sensory organ, 4—anterior sensory lobe, 5—stomal sense organ, 6—
lateral sensory organ; (C) anterior prothoracic spiracle; (D) lateral spiracular complex, spiracle to left, (E)
posterior spiracular plate; (F) compound sensilla.
several rows of acanthae circumscribe the gan. Mouth hooks bidentate (Fig. 3B-1),
body at intersegmental lines (Fig. 3A). median oral lobe rounded anteriorly, fleshy
Gnathocephalon conical, smooth, except for ventral lobe attached basally to the labial
rugose pads surrounding stomal sense or- lobe (Fig. 3B-2). Dorsal sensory organ
422 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
comprised of single papilla (Fig. 3B-3), an-
terior sensory lobe with four sensory organs
(Fig. 3B-4), stomal sense organ with three
to five sensilla (Fig. 3B-5), lateral sensillum
with single verruciform papilla (Fig. 3B-6).
Prothorax smooth, anterior thoracic spiracle
with three to four papillae (Fig. 3C). Lateral
spiracular complex on mesothorax with one
spiracle and two verruciform sensilla; meta-
thorax and abdominal segments I—VII with
one spiracle and one verruciform sensillum
(Fig. 3D). Posterior spiracular plate with
three rimae and four, single or multi-
branched, interspiracular processes (Fig.
3E). Compound sensilla comprised of two
stelex sensilla (Fig. 3F).
Third instar.—White, oblong-elliptical,
bluntly rounded on both ends, acanthae re-
stricted to area around intersegmental lines
(Fig. 4A). Mouth hooks bidentate (Fig.
4B-1), median oral lobe with numerous
fleshy protuberances on ventral lobe, at-
tached basally to labial lobe (Fig. 4B-2).
Gnathocephalon smooth, with rugose pads
laterally and ventrally near mouth lumen
(Fig. 4B). Dorsal sensory organ with a sin-
gle papilla (Fig. 4C-1), anterior sensory
lobe with four sensory organs (Fig.
4C-black lines), stomal sense organ invest-
ed with sensoria (Fig. 4C-2). Prothorax
smooth, anterior thoracic spiracle with three
to four non-protruding papillae, each with
a single slit, ecdysial scar visible (Fig. 4D).
Lateral spiracular complex with single spi-
racle and verruciform sensillum (Fig. 4B).
Posterior spiracular plate with three rimae,
and reduced interspiracular processes (Fig.
4F). Compound sensilla nearly contiguous,
each with two reduced stelex sensilla.
Puparium.—Shiny; dark fuscous to
black, some with lighter brown around the
middle; elongate-reinform, bluntly rounded
on both ends, 4.0 + 0.1 (3.1-4.3, n = 9)
mm long and 2.0 + 0.1 (1.7—2.2, n = 8)
mm wide (Fig. 5A). Anterior end with in-
vagination scar and anterior thoracic spira-
cle (Fig. 5B); posterior end with spiracular
plates non-protruding (Fig. 5C).
Diagnosis.—The immature stages of Aci-
urina thoracica Curran (Headrick and Goe-
den 1993), A. ferruginea (Doane) and A.
michaeli Goeden (Goeden and Teerink
1996a), A. idahoensis Steyskal (Goeden
and Teerink 1996b), and A. semilucida
(Bates) (Goeden and Teerink 1996c) have
been described in detail. Aciurina trixa is
the last, described California species of Aci-
urina to be examined by us. The egg of A.
trixa is smooth, nearly parallel-sided, with
a pedicel ca. 0.015 mm in length. The eggs
of A. ferruginea, A. michaeli, A. idahoensis
and A. semilucida have elongate pedicels;
the pedicel of the latter being ca. half as
long as the other species with elongate ped-
icels. The egg of A. thoracica has a short
pedicel and heavy reticulation on the pedi-
cel end that diminishes to a smooth surface
near the midpoint of the egg body.
The third instars of A. michaeli and A.
semilucida have abdominal segments III—V
expanded dorsally producing an ‘‘ambula-
tory hump.”’ No other Aciurina species in
California have this feature. The integu-
ment of A. trixa has minute acanthae re-
stricted to the areas around the interseg-
mental lines; however, A. thoracica has
acanthae in patches between the interseg-
mental lines. Aciurina idahoensis, A. sem-
ilucida and A. ferruginea have acanthae re-
stricted to the ventral surface, and A. mi-
chaeli has acanthae ventrally and laterally.
The number of sensilla associated with
the lateral spiracular complexes on the
meso- and metathorax and abdominal seg-
ments I-VII (= sensilla formula) differs
among the Aciurina species studied. The
sensilla formula, i.e. the number of lateral
spiracular sensilla on the mesothorax, the
metathorax, and the abdominal segments (a
‘?’ indicates we were unable to locate the
sensilla), for A. michaeli is 1, 1, 0; for A.
ferruginea is 3, 3, ?; for A. thoracica is ?,
1, 1; for A. idahoensis is ?, 1, 0; for A.
semilucida is 2, 2, 1; and for A. trixa is 2,
1, 1. The anterior prothoracic spiracles dif-
fer in the number of papillae among the
species studied. A. michaeli has 3—4, A. fer-
ruginea has 5, A. thoracica has 4, A. ida-
VOLUME 99, NUMBER 3
Bes
iSKU: K1000 > age
1I5KU 8720 .- @H18
Fig. 4. Third instar of A. trixa. (A) Habitus, anterior to left; (B) gnathocephalon, anterior view, 1—mouth-
hook, 2—median oral lobe; (C) sensory structures of the gnathocephalon, 1—dorsal sensory organ, 2—stomal
sensory organ; (D) anterior prothoracic spiracle; (E) lateral spiracular complex, spiracle on left; (F) caudal
segment with posterior spiracular plates.
424
“eaa9 ~1o0.oU
C.
Fig. 5. Puparium of A. trixa. (A) Habitus, anterior
to left; (B) anterior invagination scar; (C) posterior spi-
racular plates.
hoensis has 3—5, A. semilucida has 3, and
A. trixa has 4 non-protruding papillae. The
first and second instars, and the puparia do
not differ significantly in their morphology
among the species studied in California.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
BIOLOGY
Egg.—tThe eggs of A. trixa were laid sin-
gly into axillary buds on the current sea-
son’s branches (Fig. 6A) soon after adult
emergence in the spring in southern Cali-
fornia (Table 1). The eggs were inserted
partially into the tissues or among and par-
allel to the bud scales leaving the pedicel-
bearing ends exposed. Eclosion occurred
within 1 week after oviposition. The em-
bryo turned 180° and the first instar exited
basally, directly into the plant tissues.
Dodson (1987) reported that A. trixa fe-
males in New Mexico also laid eggs singly
in axillary buds of C. nauseosus with galls
becoming visible in August. Fernandez and
Price (1994) reported that oviposition oc-
curred in the field between late April and
mid-August in northern Arizona and that
eggs were laid singly in axillary buds.
First instar—The first instar tunnelled
into the axillary bud tissues and continued
down the vascular trace into the branch ba-
sipetally in the phloem between the epi-
dermis and cambium. The feeding of the
first instar stimulated the phloem and epi-
dermis outside of the woody vascular cyl-
inder surrounding the pith to expand, which
resulted in the branch swelling at and near
the axillary bud. Branch swelling was rapid
and ceased when the first larval stadium
ended. Branch swelling was also reported
by Fernandez and Price (1994). Branch tun-
nel feeding concurrent with gall growth was
also reported by Dodson (1987) and Fer-
nandez and Price (1994).
The bud scales remained as small swell-
ings at the base of galls. Gall growth con-
tinued through the spring and summer
(April—September, Table 1) as the first in-
star excavated its tunnel within the branch
from the base of the axillary bud basipetal-
ly. Thus, gall growth appeared to begin and
proceed even though no larval feeding took
place directly within the gall. Branch tun-
nels containing first instars averaged 3.1 +
0.3 (range, 2.7—3.6; n = 3) mm long. Galls
associated with branch tunnels containing
VOLUME 99, NUMBER 3 425
Fig. 6. Life stages of A. trixa. (A) egg inserted between leaf primordia (arrow); (B) mature gall, large,
smooth, pearly, gall-type; (C) second instar feeding tunnel (arrows indicate beginning and end of tunnel) and
partially excavated gall cavity; (D) second instar feeding in gall cavity and branch tunnel; (E) third instar in
gall with exit tunnel formed apically and branch tunnel packed with frass; (F) second instars in small type of
galls; (G) third instar in small gall-type; (H) puparium within gall locule, with anterior end facing exit window;
(I) adult female of A. trixa. Bars indicate ca. 1 mm.
426 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Phenological development of Aciurina trixa on Chrysothamnus nauseosus in southern California.
i—first instar; 1i—second instar; 11i—third instar.
Month
Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb
eggs eggs
i i i i i i 1
il il ll il ii
iil ill lil lil
puparium
adult adult
first instars measured 3.3 + 0.2 (range, 2.8—
3.6; n = 6) mm in diameter. First instars
confined most of their feeding to the branch
tunnel, as was also reported by Fernandez
and Price (1994). Although galls were near-
ly full-size by the end of the first stadium
in September or October (Table 1, Fig. 6B),
gall locule excavation was begun only by
the late first instars. This too, is consistent
with the findings of Dodson (1987) and
Fernandez and Price (1994). Gall locules
averaged 0.96 + 0.03 (range, 0.9-1.0; n =
3) mm wide.
Second instar.—By late October/early
November gall growth was complete (Table
1). Dodson (1987) and Fernandez and Price
(1994) also reported that gall growth was
complete by November in New Mexico and
Arizona, respectively. Galls containing sec-
ond instars averaged 3.4 + 0.07 (range,
2.3—4.1; n = 31) mm in diameter. The sec-
ond instar continued to feed within the
branch tunnel, enlarging it and extending its
length basipetally (Fig 6C). Branch tunnels
containing second instars averaged 3.3 +
0.15 (range, 2.0—6.0; n = 28) mm long. The
second instar also moved from the branch
tunnel into the fully-formed gall and began
excavating the gall lumen (Fig. 6D). The
swelling around the branch tunnel did not
increase with the continued feeding of the
second instar.
As the gall lumen was enlarged, the inner
walls became smooth; however, a substan-
tial amount of parenchymatous tissue re-
mained. Lumens of 28 galls containing sec-
ond instars averaged 1.3 + 0.08 (range,
0.5—2.5) mm wide and 3.16 + 0.3 (range,
1.6—4.3) mm long. Frass accumulated
within the gall lumen and in the branch tun-
nel as feeding continued. Frass was moved
into the tunnel from the gall and packed
into the basal end of the tunnel. The cast
exuviae and cephalopharyngeal skeletons
of first instars also were found within the
packed frass. One late-second instar was
observed beginning to form the exit tunnel
in the gall, evidenced as a narrow tunnel
toward the apex of the gall.
Third instar—Larval growth proceeded
through the fall, and by December most
galls contained third instars (Table 1). Sev-
enty galls containing third instars averaged
3.8 + 0.06 (range, 2.6—5.0) mm in diameter.
After the second molt, the third instars con-
fined most of their feeding inside the gall,
and the branch tunnel was used for frass
storage (Fig. 6E). The gall walls were re-
duced in thickness by the feeding of the
third instar. Lumens of 65 galls containing
third instars averaged 2.3 + 0.07 (range,
1.1—3.7) mm wide and 3.8 + 0.07 (range,
2.5—5.4) mm long.
Third instars completed the exit tunnel
by excavating up to the epidermis to form
a circular window for adult emergence (Fig.
6E). The formation of exit tunnels by third
instars was also reported by Dodson (1987)
and Fernandez and Price (1994). Most win-
dows were formed at or near the apices of
galls.
Larval development and feeding, gall
formation and growth, and phenological de-
velopment were similar for both gall types
VOLUME 99, NUMBER 3
occurring in southern California (Fig. 6F
G). The smaller gall type has not been re-
ported to occur outside of southern Califor-
nia (Dodson 1987, Fernandez and Price
1994).
Puparium.—Puparia were first observed
beginning in mid-February and the puparial
stage lasted ca. 2 weeks (Table 1). The pu-
parium was formed in the lumen of the gall
(Fig. 6H); no puparium was observed in a
branch tunnel. The cephalic end of the pu-
parium was oriented toward the window.
Adult.—Adults (Fig 61) are circumnatal
(Headrick and Goeden 1994) and emerged
in early spring (March—April, Table 1) fol-
lowing winter rainfall. Males were protan-
drous and awaited female emergence by
perching near galls. Males lived an average
of 32.1 + 4.0 (range, 3-57; n = 18) days;
females lived an average of 28.0 + 5.3
(range, 13-43; n = 6) days. Courtship and
mating behavior will be described and an-
alyzed in a separate paper.
Females are proovigenic and emerge
with a full complement of eggs. Mating was
observed in the field when females were
still teneral and was repeated throughout
their lifetimes. Oviposition behavior was
distinctive. Females walked or flew to the
base of a branch and then walked towards
its apex. At or near the top of the branch,
they then turned and walked down the
branch. At the first axillary bud, they
stopped and probed with the oviscape bent
downward and extruded the aculeus. If the
site was suitable, a female oviposited a sin-
gle egg after extending the aculeus into the
bud, either piercing the bud tissues or in-
serting the egg between the leaf primordia.
After oviposition, females groomed, then
moved downward to the next axillary bud
and repeated this procedure. Females did
not lay eggs into every axillary bud probed.
Females continued basipetally in this man-
ner until the base of the branch was
reached, and then either walked or flew to
another branch. Males stalked ovipositing
females and attempted and usually success-
427
fully mated with them at any point during
the above oviposition sequence.
Natural enemies.—The following were
reared from galls as primary, larval-pupal,
solitary endoparasitoids: Halticoptera sp.
(Pteromalidae), Eurytoma sp. (Eurytomi-
dae), Torymus sp. (Torymidae), Aprostoce-
tus sp., (Eulophidae), Brasema sp. (Eupel-
midae) and undetermined species of Platy-
gastridae and Petromalinae. A gregarious
species of Pronotalia sp. (Eulophidae) was
reared as a primary, larval-pupal, endopar-
asitoid from galls of A. trixa. Some mor-
tality observed in the field was attributed to
bird predation on the mature galls.
ACKNOWLEDGMENTS
We thank A. C. Sanders for identification
of plant species, Harry Andersen (Emeritus,
UC Riverside) for identification of Pter-
omalidae, John LaSalle (British Museum)
for identification of Torymidae, and Mi-
chael Gates (UC Riverside) for identifica-
tion of Eulophidae, Eupelmidae, and Pter-
omalidae. We thank EF L. Blanc, Al No-
rrbom, and Gary Steck for reviews of early
drafts of this manuscript.
LITERATURE CITED
Bates, M. 1935. Notes on American Trypetidae (Dip-
tera). II]. The genus Tephrella. Pan-Pacific Ento-
mologist 11: 103-114.
Curran, C. H. 1932. New species of Trypaneidae, with
key to the North American genera. American Mu-
seum Novitates 556: 1-19.
Dodson, G. 1987. Biological Observations on Aciurt-
na trixa and Valentibulla dodsoni (Diptera: Te-
phritidae) in New Mexico. Annals of the Ento-
mological Society of America 80: 494—500.
Dodson, G. and S. B. George. 1986. Examination of
two morphs of gall-forming Aciurina (Diptera: Te-
phritidae): ecological and genetic evidence for
species. Biological Journal of the Linnean Society
29: 63-79.
Fernandez, G. W. and P. W. Price. 1994. Life history,
courtship, and mating behavior of the gall-forming
Aciurina trixa (Diptera: Tephritidae) on Chryso-
thamnus nauseosus hololeucus (Asteraceae). Pro-
ceedings of the Entomological Society of Wash-
ington 96: 301-307.
Foote, R. H. and FE L. Blanc. 1963. The Fruit Flies or
Tephritidae of California. Bulletin of the Califor-
nia Insect Survey 7: 1-117.
428
Foote, R. H., E L. Blanc, and A. L. Norrbom. 1993.
Handbook of the Fruit Flies (Diptera: Tephritidae)
of America North of Mexico. Cornell University
Press, Ithaca.
Goeden, R. D. and J. A. Teerink. 1996a. Life histories
and descriptions of adults and immature stages of
two cryptic species Aciurina ferruginea (Doane)
and A. michaeli new species (Diptera: Tephriti-
dae), on Chrysothamnus viscidiflorus (Hooker)
Nuttall in southern California. Proceedings of the
Entomological Society of Washington 98: 415—
438.
. 1996b. Life history and descriptions of adults
and immature stages of Aciurina idahoensis
Steyskal (Diptera: Tephritidae) on Chrysothamnus
viscidiflorus (Hooker) Nuttall in southern Califor-
nia. Proceedings of the Entomological Society of
Washington 98: 681-694.
. 1996c. Life history and descriptions of adults
and immature stages of Aciurina semilucida
(Bates) (Diptera: Tephritidae) on Chrysothamnus
viscidiflorus (Hooker) Nuttall in southern Califor-
nia. Proceedings of the Entomological Society of
Washington 98: 752-766.
Headrick, D. H., and R. D. Goeden. 1991. Life history
of Trupanea californica Malloch (Diptera: Te-
phritidae) on Gnaphalium spp. in southern Cali-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
fornia. Proceedings Entomological Society Wash-
ington 93: 559-570.
. 1993. Life history and description of imma-
ture stages of Aciurina thoracica Curran (Diptera:
Tephritidae) on Baccharis sarothroides Gray in
southern California. Annals of the Entomological
Society of America 86: 68-79.
1994. Reproductive behavior of California
fruit flies and the classification and evolution of
Tephritidae (Diptera) mating systems. Studia Dip-
terologica 1: 194-252.
Munz, C. A. 1968. Supplement to A California Flora.
University of California Press, Berkeley.
1974. A Flora of Southern California. Uni-
versity of California Press, Berkeley.
Munz, C. A. and D. D. Keck. 1959. A California Flo-
ra. University of California Press, Berkeley.
Norrbom, A. L. 1989. The status of Urophora acuti-
cornis and U. sabroskyi (Diptera: Tephritidae).
Entomological News 100: 59-66.
Steyskal, G. C. 1984. A synoptic revision of the genus
Aciurina Curran, 1932 (Diptera: Tephritidae). Pro-
ceedings of the Entomological Society of Wash-
ington 86: 582-598.
Wangberg, J. K. 1981. Gall-forming habits of Aciuri-
na species (Diptera: Tephritidae) on rabbitbrush
(Compositae: Chrysothamnus spp.) in Idaho. Jour-
nal of the Kansas Entomological Society 54: 711—
W32.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 429-439
A NEW GENUS AND REDESCRIPTIONS FOR AFRICAN SPECIES
PREVIOUSLY PLACED IN ACENTRELLA (EPHEMEROPTERA: BAETIDAE)
C. R. LUGo-OrRTIZ AND W. P. MCCAFFERTY
Department of Entomology, Purdue University, West Lafayette, IN 47907, U.S.A.
Abstract.—Within the Ephemeroptera family Baetidae, Demoreptus, new genus, is
erected to accommodate D. capensis (Barnard), new combination, D. monticola (Crass),
new combination, and D. natalensis (Crass), new combination. These species were orig-
inally described from southern Africa and assigned to Acentrella Bengtsson. Demoreptus
is distinguished from Acentrella in the larval stage by having apicolaterally prolonged
and attenuated fused incisors and relatively long, well-demarcated labial palp segments
and in the adult stage by having a small, rounded process anteriorly on the mesoscutum.
The larval stage of each species in Demoreptus is redescribed and a key to the species is
provided. Analysis of interspecific relationships indicates that D. capensis is the most
plesiotypic species, and that D. natalensis and D. monticola represent derived sister spe-
cies.
Key Words:
rica
The Baetis complex (Ephemeroptera:
Baetidae) is a monophyletic grouping of
genera comprised of Acentrella Bengtsson,
Baetiella Uéno, Baetis Leach, Barbaetis
Waltz and McCafferty, Cymulabaetis
McCafferty and Waltz, Gratia Thomas,
Heterocloeon McDunnough, Labiobaetis
Novikova and Kluge, Liebebiella Waltz and
McCafferty, Platybaetis Miiller-Liebenau,
and Tanzaniella Gillies. The complex is de-
fined by the possession of the villopore, lo-
cated on the ventral margin near the base
of the larval femora (see, e.g. Waltz and
McCafferty 1987: Figs. 1, 4, 5, 12, 17). Pre-
cise phylogenetic relationships within the
complex, however, remain unknown.
The Baetis complex is relatively common
and diverse in the Holarctic and Orient, but
it is poorly known in the Neotropics and
Australia. In the Afrotropics, the genera
Acentrella, Baetis, Pseudocloeon, and Tan-
zaniella have been variously treated by Bar-
Ephemeroptera, Baetidae, Demoreptus, new genus, new combinations, Af-
nard (1932, 1940), Crass (1947), Kimmins
(1955, 1960), Demoulin (1970), Kopelke
(1980), and Gillies (1991, 1993, 1994).
Only one species has been described in the
poorly known genus Tanzaniella (Gillies
1991), and, as Waltz and McCafferty (1987)
and McCafferty and de Moor (1995) point-
ed out, the taxonomic status of species as-
signed to Acentrella, Baetis, and Pseudo-
cloeon are in need of re-evaluation. This
situation remains the same even though Gil-
lies (1994) transferred all African species of
Pseudocloeon to Baetis.
In this study, we address the African spe-
cies that were described in the genus Acen-
trella: A. capensis Barnard, A. monticola
Crass, and A. natalensis Crass. Those spe-
cies proved to be referable to a new genus
described herein. We additionally redescri-
be the larval stage of each species and pro-
vide a key for their identification. Materials
examined are housed in the Albany Muse-
430 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
um (AM), Grahamstown, Eastern Cape
Province, South Africa, and the Purdue En-
tomological Research Collection (PERC),
West Lafayette, Indiana, U.S.A.
Demoreptus Lugo-Ortiz and
McCafferty, new genus
Larva (Fig. 1).—Head: Antennae 1.0—
2.0 length of head capsule. Labrum with
deep (Figs. 3, 16) or shallow (Fig. 9) an-
teromedial notch. Mandibles with apicola-
terally prolonged and attenuated fused in-
cisors, with distinct but poorly developed
denticulation (Figs. 4, 5, 18, 19) or adenti-
culate and bladelike (Figs. 11, 12). Maxillae
(Figs. 6, 13, 20) with palps two segmented.
Labium (Figs. 7, 14, 21) with relatively
long glossae and paraglossae; palps rela-
tively long, extending beyond apices of
glossae and paraglossae; palp segment 2
with poorly defined distomedial lobe; palp
segment 3 width not exceeding that of api-
cal width of segment 2 (Figs. 7, 21) or me-
dially broader than apical width of segment
2 (Fig. 14). Thorax: Hindwingpads rudi-
mentary to well developed. Legs with vil-
lopore present; femora with long, fine, sim-
ple setae dorsally; tarsi twisted; tarsal claws
with single row of denticles and two sub-
apical short, fine, simple setae. Abdomen:
Terga without scales or scale bases; poste-
rior marginal spines poorly defined. Gills
on abdominal segments 1-7, platelike,
poorly tracheated, marginally smooth (Fig.
23). Paraprocts (Figs. 8, 15, 24) without
marginal spines. Two caudal filaments pres-
ent, with abundant setae medially; medial
caudal filament reduced to one segment.
Adult.—Mesoscutum with small, round-
ed anterior process in lateral view (Fig. 2).
Forewings with paired marginal intercalar-
ies. Hindwings present, except in males of
D. natalensis; when present, with (Barnard
1932: Fig. 12b) or without (Crass 1947:
Fig. 16b, 18b) costal process. Male genital
forceps (Crass 1947: Fig. 17h) three seg-
mented; segment 2 strongly bowed inward-
ly; segment 3 ellipsoidal.
Etymology.—We are honored in naming
this genus after Ferdinand C. de Moor, for
his support of the study of aquatic insects
in South Africa. The name consists of an
arbitrary combination of letters that incor-
porates a Latinization of our colleague’s last
name and the Latin suffix reptus (to crawl),
which is an allusion to the behavior of the
larvae.
Type species.—Demoreptus natalensis
(Crass), 1947: 72.
Species included.—Demoreptus capensis
(Barnard), new combination; D. monticola
(Crass), new combination; D. natalensis
(Crass), new combination.
Distribution.—Lesotho; South Africa:
Eastern Cape, KwaZulu-Natal, Mpumalan-
ga, Western Cape.
Discussion.—Outgroup comparisons
with Baetis indicate that Demoreptus prob-
ably arose from a Baetis-like ancestor
whose adults had hindwings with a costal
process and whose larvae possessed a la-
bium with relatively long glossae, paraglos-
sae, and palp segments; relatively short,
ventrally oriented legs with straight tarsi;
well-developed hindwingpads; terga with
scales and well-developed posterior mar-
ginal spines; marginally serrate gills; and a
well-developed medial caudal filament. De-
moreptus shows the following apomor-
phies: apicolaterally prolonged and attenu-
ated fused incisors; relatively long, out-
stretched legs with twisted tarsi; loss of ter-
gal scales; reduction of tergal posterior
marginal spines; loss of marginal serrations
in the gills; and reduction of the medial
caudal filament.
Demoreptus is phenotypically somewhat
similar to the Holarctic and Oriental genus
Acentrella as redefined by Waltz and
McCafferty (1987). Larvae of both genera
lack tergal scales and marginal spines on
the gills and have outstretched legs with
twisted tarsi, reduced marginal spines on
the abdominal terga, and a reduced medial
caudal filament. However, larvae of De-
moreptus have relatively long labial palps
(especially with regard to segment 2) with
segments that are well demarcated (Figs. 7,
431
VOLUME 99, NUMBER 3
larva (dorsal).
Demoreptus capensis,
Fig. 1.
432 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
6
Figs. 2-8.
8
Demoreptus capensis. 2, Mesoscutum (lateral; pointer towards process). 3, Labrum (dorsal). 4,
Left mandible. 5, Right mandible. 6, Right maxilla. 7, Labium (left-ventral; right-dorsal). 8, Paraproct.
14, 21). More importantly, adults of De-
moreptus have a small, rounded process an-
teriorly on the mesoscutum (Fig. 2) that is
distinctly different from the apomorphically
pointed, dorsally oriented process that
adults of Acentrella and Heterocloeon have
(Waltz 1994: Fig. 1). Adults of the poorly
known Afrotropical genus Tanzaniella
probably have that process (R. D. Waltz,
pers. comm.), and it is possible that adults
of the Oriental genus Liebebiella will also
prove to have it once they are known (Waltz
1996). In any case, the presence of a point-
ed, dorsally oriented process appears to in-
dicate a monophyletic grouping within the
Baetis complex, and because Demoreptus
lacks that process, it cannot be considered
to belong in that grouping.
Larvae of Demoreptus, Acentrella, and
certain other species of the Baetis complex,
as well as distantly related genera such as
the Afrotropical genus Acanthiops Waltz
and McCafferty and the Panamerican genus
Baetodes Needham and Murphy, show sev-
VOLUME 99, NUMBER 3
eral convergent adaptations for sprawling
and clinging on rocks in high gradient, fast-
running streams. Those adaptations include
long, outstretched legs, relatively small
gills, and reduced medial caudal filament
(see, e.g. Lugo-Ortiz and McCafferty 1996
and Barber-James and McCafferty (1997),
and in some instances the taxa that have
them look strikingly similar and may be
easily confused, such as is the case with
Demoreptus and Acentrella.
Demoreptus capensis (Barnard),
new combination
(Figs. 1-8)
Acentrella capensis Barnard 1932: 259 (lar-
va; male, female adults).
Baetis capensis (Barnard): Demoulin 1970:
66.
Larva (Fig. 1).—Body length: 6.5—7.0
mm; caudal filaments length: 3.8—4.0 mm.
Head: Coloration yellow-brown, with no
distinct pattern. Antennae 2.0 length of
head capsule. Labrum (Fig. 3) nearly 1.22
wider than long, deeply cleft anteromedi-
ally, with submedial long, fine, simple seta
and five to six long, fine, simple setae on
either side of midline. Hypopharynx similar
to Figure 17. Left mandible (Fig. 4) with
six denticles; marginal lateral denticle en-
larged. Right mandible (Fig. 5) with six
denticles; marginal lateral denticle en-
larged. Maxillae (Fig. 6) with four denticles
on galealaciniae and five to six fine, simple
setae on medial hump. Labium (Fig. 7) with
glossae and paraglossae subequal in length;
paraglossae somewhat broad; palp segment
1 as long as segments 2 and 3 combined;
palps segment 2 with five to six fine, simple
setae dorsally; palp segment 3 more or less
apically rounded, not bulbous or clublike
(width not exceeding that of apical width of
segment 2). Thorax: Coloration yellow-
brown, with irregular medium brown mark-
ings. Hindwingpads long, almost reaching
hind margin of abdominal segment 1. Legs
yellow-brown; femora with poorly devel-
oped villopore, row of long, robust, simple
433
setae dorsally, and scattered short, stout and
short, fine, simple setae ventrally; tibiae
with row of short, fine, simple setae dor-
sally and scattered short, stout, and short,
fine, simple setae ventrally; tarsi with scat-
tered short, fine, simple setae dorsally and
six to seven stout, simple setae increasing
in length towards distal end ventrally; tarsal
claws with 11-12 denticles, increasing in
length and girth distally. Abdomen: Color-
ation yellow-brown, with medium and dark
brown markings. Segment | yellow-brown;
segments 2—6 yellow-brown, with pair of
dark brown longitudinal submedial mark-
ings and dark brown hind margins; seg-
ments 7—9 with pair of oblique medium
brown dashes anteriorly, pair of small
specks in posterior half, and medium brown
to dark brown hind margins; segment 10
yellow-brown, with diffuse markings. Ster-
na pale yellow-brown. Gills whitish, poorly
tracheated. Paraprocts as in Figure 8. Cau-
dal filaments cream to pale yellow.
Adult.—See description of Barnard
(1932):
Material examined.—S5 larvae, LESO-
THO, Mokhotlong-Sengu, basin below
Woolshed on Schonghong R, IX-25-1988,
P. H. Skelton (AM); 2 larvae, LESOTHO,
Sani-Linakeng Basin at road drift, tributary
of Sani R, [X-24-1988, P H. Skelton (AM);
3 larvae, SOUTH AFRICA, Eastern Cape,
Waterkloof, HI-27-1991, E Weir (AM); 2
larvae, SOUTH AFRICA, KwaZulu-Natal,
Highmoor Forest, Littlke Mooi R, 1800 m,
IX-19-1990, W. P. and N. McCafferty
(PERC); 2 larvae, SOUTH AFRICA, West-
ern Cape, Jonkershoek Mnts., 2nd tributary
of Eerste R, 1000 m, IX-28-1990, W. P. and
N. McCafferty (PERC); 33 larvae, 1
adult, SOUTH AFRICA, Western Cape,
Jonkershoek Mnts., Eerste R nr bridge at
end of dirt rd, [X-28-1990, W. P. and N.
McCafferty (PERC).
Discussion.—Larvae of D. capensis are
distinguished from those of D. natalensis
by the setation of the labrum (Fig. 3), the
deeper and sharper denticulation of both
mandibles (Figs. 4, 5), relatively short seg-
434 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ment 2 and apically rounded segment 3 of
the labial palps (Fig. 7), and long hind-
wingpads that almost reach the hind margin
of abdominal segment 1. Adults of D. ca-
pensis are distinguished from those of D.
natalensis by the presence of an acute cos-
tal process in the hindwings (Barnard 1932,
1940, Crass 1947).
Demoreptus monticola (Crass),
new combination
(Figs. 9-15)
Acentrella monticola Crass 1947: 75 (larva;
male, female adults).
Baetis monticola (Crass): Demoulin 1970:
68.
Larva.—Body length: 6.5—6.8 mm; cau-
dal filaments length: 5.8-6.0 mm. Head:
Coloration yellow-brown, with vermiform
medium brown markings on frons. Antenna
1.5X length of head capsule. Labrum (Fig.
9), 2.62 wider than long, with shallow an-
teromedial emargination and 18—20 long,
fine, simple setae on either side of midline;
submedial seta absent. Hypopharynx as in
Figure 10. Left mandible (Fig. 11) with in-
cisors fused into one bladelike structure
lacking denticles; prostheca apically dentic-
ulate. Right mandible (Fig. 12) with inci-
sors fused into bladelike structure lacking
denticles; prostheca apically pointed, with
long, fine, simple setae medially. Maxillae
(Fig. 13) with four denticles on galealaci-
niae and six to seven fine, simple setae on
medial hump. Labium (Fig. 14) with glos-
sae and paraglossae somewhat slender and
subequal in length; palp segment 1 sub-
equal to segments 2 and 3 combined; palp
segment 2 with three to four fine, simple
setae dorsally; palp segment 3 bulbous,
clublike (medially broader than apical
width of segment 2). Thorax: Coloration
yellow-brown, with irregular medium
brown markings. Legs yellow-brown; fem-
ora with poorly developed villopore, row of
long, robust, simple setae dorsally, and
scattered short, stout and short, fine, simple
setae ventrally; tibiae with scattered short,
fine, simple setae dorsally and scattered
short, stout, and short, fine, simple setae
ventrally; tarsi with scattered short, fine,
simple setae dorsally and six to seven stout,
simple setae increasing in length towards
distal end ventrally; tarsal claws with 11-—
12 denticles, increasing in length and girth
distally. Abdomen: Coloration yellow-
brown and dark brown. Segment | and 2
dark brown with large yellow-brown sub-
lateral circular to oblong markings; seg-
ments 3—6 as segments | and 2, except with
slender dorsal longitudinal medial yellow-
brown streak; segments 7—9 yellow-brown,
with submedial pair of dark brown specks
in midregion and dark brown hind margins;
segment 10 yellow-brown. Sterna cream to
pale yellow-brown. Gills whitish, with sin-
gle conspicuous tracheal trunk medially.
Paraprocts as in Figure 15. Caudal filaments
cream to pale yellow.
Adult.—See description of Crass (1947).
Material examined.—8 larvae, SOUTH
AFRICA, Eastern Cape, Hogsback, Ma-
donna and Child waterfall, X-7-1989
(AM); 142 larvae, SOUTH AFRICA,
KwaZulu-Natal, Sani Pass Rd at police
post, 1950 m, 16.4°C, X-1-1971, G. EF and
C. H. Edmunds (PERC); 8 larvae,
SOUTH AFRICA, KwaZulu-Natal, Umzi-
mkulu R, between Underberg and Boes-
mansnek, 15.5°C, X-2-1971, G. F and C.
H. Edmunds (PERC); 5 larvae and 2 sub-
imago, SOUTH AFRICA, KwaZulu-Na-
tal, Pietermaritzburg Nat. Bot. Garden,
Dorspruit, I[X-18-1990, W. P. and N.
McCafferty (PERC); 31 larvae, SOUTH
AFRICA, KwaZulu-Natal, Highmoor
For., Little Mooi R, 1800 m, [X-19-1990,
W. P. and N. McCafferty (PERC); 8 lar-
vae, SOUTH AFRICA, Mpumalanga, 5
mi NE of Machadodorp, I[X-22-1971, G.
F and C. H. Edmunds and H. J. Schoon-
bee (PERC).
Discussion.—Larvae of C. monticola
are distinguished by the setation and shal-
low anterior emargination of the labrum
(Fig. 9), adenticulate and bladelike man-
dibular incisors (Figs. 11, 12), and bul-
ee
VOLUME 99, NUMBER 3 435
13 15
Figs. 9-15. Demoreptus monticola. 9, Labrum (dorsal). 10, Hypopharynx. 11, Left mandible. 12, Right
mandible. 13, Right maxilla. 14, Labium (left-ventral; right-dorsal). 15, Paraproct.
bous segment 3 of the labial palps (Fig. Demoreptus natalensis (Crass),
14). Adults of D monticola differ from new combination
those of D. capensis in lacking an acute (Figs. 16—24)
costal process in the hindwings, but evi- Acentrella natalensis Crass 1947: 72 (larva;
dently can only be told from those of D. male, female adults).
natalensis by the presence of hindwings Baetis natalensis (Crass): Demoulin 1970:
in both sexes (Crass 1947). 68.
436 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
20
24
Figs. 16-24. Demoreptus natalensis. 16, Labrum (dorsal). 17, Hypopharynx. 18, Left mandible. 19, Right
mandible. 20, Right maxilla. 21, Labium (left-ventral; right-dorsal). 22, Gill 4. 23, Detail of gill margin. 24,
Paraproct.
Larva.—Body length: 4.8—5.1 mm; cau-
dal filaments length: 4.8-5.1 mm. Head:
Coloration dark brown, with vermiform
yellow-brown markings on frons. Antennae
as long as head capsule. Labrum (Fig. 16)
nearly 1.81 wider than long, anteromedi-
ally deeply cleft, with submedial long, fine,
simple seta and 10—11 long, fine, simple se-
tae on either side of midline. Hypopharynx
as in Figure 17. Left mandible (Fig. 18)
with six denticles; marginal lateral denticle
small. Right mandible (Fig. 19) with six
denticles; marginal lateral denticles small.
Maxillae (Fig. 20) with four denticles on
galealaciniae and five to six fine, simple se-
tae on medial hump. Labium (Fig. 21) with
VOLUME 99, NUMBER 3
glossae subequal to paraglossae; paraglos-
sae somewhat broad and long; palp segment
1 subequal to segments 2 and 3 combined;
segment 2 with three to four fine, simple
setae dorsally; palp segment 3 more or less
pointed apically, not bulbous or clublike
(width not exceeding that of apical width of
segment 2). Thorax: Coloration dark
brown, with irregular dark yellow-brown
markings. Hindwingpads short, not more
than 0.25 length of abdominal segment 1.
Legs medium brown with irregular yellow-
brown markings; femora with poorly de-
veloped villopore, row of long, robust, sim-
ple setae dorsally, and scattered short, stout
and short, fine, simple setae ventrally; tibiae
with row of short, fine, simple setae dor-
sally and scattered short, stout, and short,
fine, simple setae ventrally; tarsi with row
of short, fine, simple setae dorsally and
eight to nine stout, simple setae increasing
in length towards distal end ventrally; tarsal
claws with 11—12 denticles, increasing in
length and girth distally. Abdomen: Color-
ation dark brown to dark yellow-brown,
with medium brown markings. Segment 1
dark yellow-brown, with dark brown ante-
rior, posterior, and lateral margins; segment
2 dark brown, somewhat paler in midre-
gion; segment 3 dark brown, with large
sublateral dark yellow-brown quadrangles;
segments 4 and 5 as segment 1, except with
anterior submedial pair of small dark brown
dashes; segments 6 and 7 as segment 1, ex-
cept with submedial medium-sized dark
brown circles in midregion; segments 8 and
9 as segments 6 and 7, except pale yellow-
brown; segment 10 pale yellow-brown.
Sterna pale yellow-brown to cream. Gills
(Fig. 22) untracheated, dark yellow-brown
to medium brown in midregion and whitish
along margin; margin (Fig. 23) smooth and
with fine, simple setae. Paraprocts as in Fig-
ure 24. Caudal filaments pale yellow-brown
to cream.
Adult.—See description of Crass (1947).
Material examined.—7 larvae, LESO-
THO, Schonghong R, Sangu-Orange Basin,
Sani Rd bridge, [X-21-1988, P. H. Skelton
437
(AM); 2 larvae, SOUTH AFRICA, Eastern
Cape, Berg R, at Hwy N2, nr Grahams-
town, XI-13-1990, W. P. and N. McCafferty
(PERC); 4 larvae, SOUTH AFRICA, East-
ern Cape, Hogsback, Buffalo R, Madonna
and Child waterfall, X-7-1989 (AM); 62
larvae, SOUTH AFRICA, KwaZulu-Natal,
Sani Pass Rd at police post, 1950 m,
16.4°C, X-1-1971, G. FE and C. H. Edmunds
(PERC); 5 larvae, SOUTH AFRICA,
KwaZulu-Natal, Howick Falls, Umgeni R,
2-X-71, G. EK and C. H. Edmunds (PERC);
18 larvae, SOUTH AFRICA, KwaZulu-Na-
tal, Impendle, W fork of Furth R, 1450 m,
IX-18-1990, W. P. and N. McCafferty
(PERC); larva, SOUTH AFRICA, Kwa-
Zulu-Natal, Impendle, Furth R, 1250 m,
IX-18-1990, W. P. and N. McCafferty
(PERC); 2 larvae, SOUTH AFRICA,
KwaZulu-Natal, Camberg Nat. Res., riffle
in Mooi R, IX-19-1990, W. P. and N.
McCafferty (PERC); 36 larvae, SOUTH
AFRICA, KwaZulu-Natal, Krantzloof Nat.
Res., Molweni stream nr Kloof, 16.4°C, G.
F and C. H. Edmunds, X-4-1971 (PERC);
11 larvae, SOUTH AFRICA, KwaZulu-Na-
tal, Molweni R at Krantzloof Nat. Res., 978
m, nr Durban, [X-21-1990, W. P. and N.
McCafferty (PERC); 21 larvae, SOUTH
AFRICA, Mpumalanga, MacMac R, above
MacMac Falls, nr. Graskop, 1820 m, W. P.
and N. McCafferty (PERC); 3 larvae,
SOUTH AFRICA, Mpumalanga, Long
Tom St. For., upper Sabie R, X-26-1990, W.
P. and N. McCafferty (PERC).
Discussion.—Larvae of D. natalensis are
distinguished from those of D. capensis by
the setation of the labrum (Fig. 16), small
marginal incisors of the mandibles (Figs.
18, 19), the apically narrowly rounded la-
bial palps (Fig. 21), and brown abdominal
gills that are submarginally whitish (Fig.
22). Adults of D. natalensis can be sepa-
rated from those of D. capensis by the ab-
sence of a costal process in the hindwings,
and evidently can be separated from those
of D. monticola by the absence of hind-
wings in its males (Crass 1947).
438
KEY TO THE LARVAE OF DEMOREPTUS
1. Labial palps segment 3 bulbous and clublike
(medially broader than apical width of segment
2) (Fig. 14); labrum with shallow anteromedial
emargination (Fig. 9); incisors of mandibles
adenticulate and bladelike (Figs. 11, 12) ...
D. monticola
— Labial palps segment 3 not bulbous, narrowly
or broadly rounded apically (width not exceed-
ing that of apical width of segment 2) (Figs. 7,
21); labrum anteromedially deeply cleft (Figs.
3, 16); incisors of mandibles with distinct den-
MeMatonn(bies 4 esos, 9) ih oe weNeu en 2
2. Labial palps segment 3 narrowly rounded api-
cally (Fig. 21); hindwingpads less than 0.25
length of abdominal segment 1; abdominal
gills brown, submarginally whitish (Fig. 22)
a ve aft ad Se Al ea Rae Cie care D. natalensis
— Labial palps segment 3 broadly rounded api-
cally (Fig. 7); hindwingpads long, almost
reaching hind margin of abdominal segment 1;
abdominal gills whitish throughout .. D. capensis
INTERSPECIFIC RELATIONSHIPS
Using Baetis as our outgroup for cladistic
analysis of species relationships within De-
moreptus, we hypothesize that D. capensis
is the most plesiotypic species, whereas D.
monticola and D. natalensis represent sister
species. Overall, however, D. capensis ap-
pears most similar to D. natalensis; both
have anteriorly notched labra with a sub-
medial pair of long, simple setae (Figs. 3,
16), denticulate mandibles (Figs. 4, 5, 18,
19), and well-demarcated labial palps with
a segment 3 whose width does not exceed
that of segment 2 (Figs. 7, 21). Those sim-
ilarities, however, are based on symple-
siomorphies, and therefore are not indica-
tive of common ancestry between the two
species. Furthermore, D. monticola is a
highly evolved species and thus very dis-
tinctive. Nevertheless, we consider the loss
of the costal process in the hindwings of D.
monticola and D. natalensis (Crass 1947:
Figs. 16b, 18b) a compelling synapomor-
phy indicative of the recent common an-
cestry of these species. Demoreptus capen-
sis retains the plesiomorphic acute costal
process (Barnard 1932: Fig. 12b) in the
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
hindwings, similar to that seen in Baetis
and most other baetines.
Because, phenotipically, D. monticola is
the most distinct of the three species and D.
capensis and D. natalensis are most similar
to each other, it would be tempting to erect
a separate genus for D. monticola. How-
ever, it is not cladistically allowable be-
cause a separate taxon for D. capensis and
D. natalensis would be paraphyletic. This
situation is similar to that among the spe-
cies of Acanthiops, where A. marlieri (De-
moulin), being highly evolved, shows an
extreme expression of clinal characters that
may lead incorrectly to interpret it as a sep-
arate clade (see Barber-James and Mc-
Cafferty 1997).
ACKNOWLEDGMENTS
We thank H. Barber-James and E de
Moor (Grahamstown, South Africa) and N.
McCafferty (West Lafayette, Indiana) for
their assistance in the field. We also thank
A. V. Provonsha (West Lafayette, Indiana)
for the larval habitus drawing. Research
funds were provided to WPM by the South
African Foundation for Research Develop-
ment and the Anglo-American and de Beers
Chairman’s Fund. The Albany Museum
kindly provided office and laboratory facil-
ities to WPM during his stay in South Af-
rica. This paper has been assigned Purdue
Agricultural Research Program Journal No.
15246.
LITERATURE CITED
Barber-James, H. and W. P. McCafferty. 1997. Review
and a new species of the African genus Acan-
thiops (Ephemeroptera: Baetidae). Annales de
Limnologie 73(2): 1-8.
Barnard, K. H. 1932. South African may-flies
(Ephemeroptera). Transactions of the Royal So-
ciety of South Africa 20: 201—259.
. 1940. Additional records and descriptions of
new species of South African alder-flies (Megal-
optera), may-flies (Ephemeroptera), caddis-flies
(Trichoptera), stone-flies (Perlaria), and dragon-
flies (Odonata). Annals of the South African Mu-
seum 32: 609-661.
Crass, R. S. 1947. The may-flies (Ephemeroptera) of
VOLUME 99, NUMBER 3
Natal and Eastern Cape. Annals of the Natal Mu-
seum 11: 37-110.
Demoulin, G. 1970. Ephemeroptera des faunes éthio-
pienne et malgache. South African Animal Life
14: 24-170.
Gillies, M. T. 1991. A diphyletic origin for the two-
tailed baetid nymphs occurring in east African
stony streams with a description of the new genus
and species Tanzaniella spinosa gen. nov. sp.
nov., pp. 175-187. In Alba-Tercedor, J., and A.
Sanchez-Ortega, eds., Overviews and strategies of
Ephemeroptera and Plecoptera. Sandhill Crane
Press, Gainesville, Fla.
1993. Descriptions of some Afrotropical
Baetidae (Ephemeroptera). II. Baetis Leach, s.l.,
West African species. Aquatic Insects 15: 213—
225;
1994. Descriptions of some Afrotropical
Baetidae (Ephemeroptera). I]. Baetis Leach, s.l.,
East African species. Aquatic Insects 16: 105—
118.
Kimmins, D. E. 1955. Ephemeroptera from Nyasa-
land, with descriptions of three new species and
some interesting nymphal forms. Annals and
Magazine of Natural History 8: 859-880.
. 1960. Notes on East African Ephemeroptera,
with descriptions of new species. Bulletin of the
439
British Museum (Natural History), Entomology 9:
339-355.
Kopelke, J.-P. 1980. Ephemeroptera aus der Emergenz
des zentralafrikanischen Bergbaches Kalengo
(Zaire). Teil 1: Baetidae. Entomologische Abhan-
dlungen, Staatliches Museum fiir Tierkunde in
Dresden 43: 99-129.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1996. Phy-
logeny and classification of the Baetodes complex
(Ephemeroptera: Baetidae) with description of a
new genus. Journal of the North American Ben-
thological Society 15: 367-380.
McCafferty, W. P. and E C. de Moor. 1995. South
African Ephemeroptera: problems and priorities,
pp. 463—476. In Corkum, L., and J. Ciborowski,
eds., Current directions on Ephemeroptera, Cana-
dian Scholars’ Press, Toronto.
Waltz, R. D. 1994. Field recognition of adult Acen-
trella and Heterocloeon (Ephemeroptera: Baeti-
dae). The Great Lakes Entomologist 26: 321-323.
. 1996. Acentrella feminalis, new combination
for an Oriental Baetis (Ephemeroptera: Baetidae).
Aquatic Insects 18: 111-112.
Waltz, R. D. and W. P. McCafferty. 1987. Systematics
of Pseudocloeon, Acentrella, Baetiella, and Lie-
bebiella new genus (Ephemeroptera: Baetidae).
Journal of the New York Entomological Society
95: 553-568.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 440-460
A NEW SPECIES OF MEALYBUG IN THE GENUS DYSMICOCCUS
(HEMIPTERA: COCCOIDEA: PSEUDOCOCCIDAE) OF IMPORTANCE IN
HIGHBUSH BLUEBERRIES (VACCINIUM CORYMBOSUM, ERICACEAE) IN
THE EASTERN UNITED STATES
DOUGLASS R. MILLER AND SRIDHAR POLAVARAPU
(DRM) Systematic Entomology Laboratory, PSI, Agricultural Research Service,
USDA, Bldg. 046, BARC-W, Beltsville, MD 20705, U.S.A.; (SP) Blueberry and Cran-
berry Research Center, Rutgers University, Chatsworth, NJ, 08019, U.S.A.
Abstract.—A new species of mealybug, Dysmicoccus vaccinii Miller and Polavarapu,
is described that is believed to be a pest of highbush blueberries, Vaccinium corymbosum
L. It has an unusual life history since most instars can be found in the field throughout
the year, including January and February. The four female instars and five male instars
are described and illustrated, including apterous and macropterous adult males. The blue-
berry mealybug is suggested as a common name. New synonymy is included as follows:
Dysmicoccus bispinosus Beardsley is considered to be a junior synonym of D. texensis
(Tinsley).
Key Words:
Mealybugs, Coccoidea, Pseudococcidae, pest, blueberry, Vaccinium, roots,
ants, Acanthomyops, Lasius, mutualism
Infestations of an undescribed species of
mealybug belonging to the genus Dysmi-
coccus were discovered several years ago
on the roots of highbush blueberries (Vac-
cinium corymbosum L.) in southern New
Jersey. This genus encompasses more than
100 species (Ben-Dov 1994) including sev-
eral major pests such as the gray sugarcane
mealybug, D. boninsis (Kuwana) and the
pineapple mealybug, D. brevipes (Cocker-
ell).
The infestations n New Jersey are be-
coming increasingly prevalent especially in
the light sandy-loam soils of Atlantic Coun-
ty. Although the impact of the mealybug on
blueberry production and fruit quality has
not been quantified, severe infestations ap-
pear to reduce vigor and lead to stunting of
young plants. Circumstantial evidence has
implicated this species as a vector of the
Red ringspot virus (belonging to the Cau-
limovirus group of viruses), the causal
agent of the Red ringspot disease in blue-
berries (Ramsdell et al. 1987). Red ringspot
is one of the most important viral diseases
of blueberries in New Jersey and also oc-
curs in Arkansas, Connecticut, Massachu-
setts, Michigan, New York, North Carolina
and Oregon (Ramsdell et al. 1987).
The purpose of this research is 1) to
name this species and describe its morpho-
logical characters so that it can be differ-
entiated from similar species, 2) to provide
preliminary information on the biology of
the species, and 3) to incorporate it in a key
to adult females of Dysmicoccus so that the
new species can be accurately identified.
In order to describe the new species and
compare it with the most similar species, it
is necessary to make the following nomen-
clatural changes effecting the status of Dys-
micoccus bispinosus Beardsley and D. tex-
VOLUME 99, NUMBER 3
ensis (Tinsley). DRM has examined type
specimens of these species and concludes
that they belong to only one species (new
synonymy). Since D. texensis was de-
scribed by Tinsley in 1900 and D. bispi-
nosus was described by Beardsley in 1965,
by the law of priority the correct name of
the species is Dysmicoccus texensis. Since
type material of Pseudococcus texensis is a
syntype series, we have selected as lecto-
type the left adult female specimen mount-
ed on a slide with | other adult female with
the left label ‘““Dactylopius/ texensis (Tin-
sley)/Type/on Acacia/ farnesiana/ San Di-
ego, Texas/ E. A. Schwarz coll./ Dec. 1895
1899”’; the right label contains a map giv-
ing the location of the lectotype and states
‘*Pseudococcus/ texensis [LECTOTY PE/
PARALECTOTYPE/ .” In addition to the
lectotype there are 28 adult female paralec-
totypes on 6 slides; all specimens are in the
USNM.
METHODS
To determine the percent of the mealybug
population in different stages at various
times of the year, 1—2 infested blueberry
plants were collected at approximately 1—2
month intervals from an infested blueberry
field in Hammonton, New Jersey. Plants
were gently uprooted with minimal distur-
bance to the root system. Each plant along
with the surrounding soil were placed in a
30-gal plastic bag for further examination.
In the laboratory, the root-system and the
accompanying soil from each plant sample
were carefully examined for different
mealybug stages. This sampling procedure
may be biased against the minute, early in-
star nymphal stages, but should neverthe-
less provide a qualitative measure of the oc-
currence of various stages of the insect.
Mealybug samples were preserved in 70%
alcohol and shipped to the Systematic En-
tomology Laboratory (SEL) for identifica-
tion of different stages of the insect.
Terminology in the descriptions follows
that of Williams and Granara de Willink
(1992) and Gimpel and Miller (1996) for
441
adult females and immatures and that of Af-
ifi (1968) for adult males. Measurements
and numbers are from 10 specimens when
available, and are given as an average fol-
lowed by the range in parentheses. Enlarg-
ments on illustrations are not proportional.
Depositories of specimens are: The Natural
History Museum, London (BMNH); Cali-
fornia Department of Food and Agriculture,
Sacramento (CDA); Florida State Collec-
tion of Arthropods, Gainesville (FSCA);
Muséum National d’ Histoire Naturelle, Par-
is (MNHN); University of California, Davis
(UCD); National Museum of Natural His-
tory, Beltsville, MD (USNM).
RESULTS
Dysmicoccus vaccinii Miller and
Polavarapu, new species
Suggested Common Name: Blueberry
mealybug
Type data.—The adult female holotype is
mounted alone on a slide with the left label
“NEW JERSEY/ Hammonton, Variety/
Farms, Atlantic Co.,/ 17-XI-1994/ ex. Blue-
berry/ S. Polavarapu”’ right label ““Dysmi-
coccus/ vaccinii/ Miller and Polavarapu/
HOLOTYPE” . This slide is deposited in
the USNM. In addition there are 1,354
paratypes on 185 slides that are deposited
in BMNH, CDA, FSCA, MNHN, UCD,
USNM.
Etymology.—The species epithet is the
genitive form of the blueberry host genus
Vaccinium.
ADULT FEMALE
(Fig. 1)
Slide-mounted characters.—Holotype
oval, length 1.9 mm, width 1.1 mm. Para-
types 1.7(1.4—-1.9) mm long, 1.1(0.8—1.2)
mm wide.
Dorsum with 17 pairs of cerarii, cerarian
formula as follows: Left side 1—6(2), 7(3),
8-11(2), 12(3), 13-14(2), 15(3), 16(2),
17(4); paratypes with cerarius 1 with 2 con-
ical setae, cerarius 2 with 2(2—3) conical se-
tae, cerarius 3 with 2(1—2) conical setae,
442
cerarius 4 with 2(2—3) conical setae, cerar-
ius 5 with 2(1—3) conical setae, cerarius 6
with 2(1—2) conical setae, cerarius 7 with
2(2—3) conical setae, cerarius 8 with 2(1-—3)
conical setae, cerarius 9 with 2(1—3) conical
setae, cerarius 10 with 3(2—3) conical setae,
cerarius 11 with 2(1—3) conical setae, cer-
arius 12 with 3(3—4) conical setae, cerarius
13 with 2(1-—3) conical setae, cerarius 14
with 2(2—3) conical setae, cerarius 15 with
3(2—5) conical setae, cerarius 16 with 5(3—
7) conical setae, cerarius 17 with 4(3—5)
conical setae. Cerarius 12 with 3 auxiliary
setae (paratypes with 3(1—5) setae), 24 tri-
locular pores (paratypes with 23(17-27)
pores), and 5 discoidal pores (paratypes
with 3(1—5) pores). Multilocular pores ab-
sent; trilocular pores evenly scattered over
surface; discoidal pores about equal to di-
ameter of trilocular pore. Oral-collar tubular
ducts absent. Longest submedial seta on
segment VII 27 pw long (paratypes 30(22—
37) 2); 6 submedial setae on segment VIII
(paratypes 6(5—6) setae), longest seta 30 w
long (paratypes 34(25—42) ).
Anal ring seta 116 w long (paratypes
117(101-143) wp); 1.4 times as long as
greatest diameter of ring (paratypes
1.4(1.3—1.6) times).
Venter with multilocular pores in poste-
rior and anterior bands on segments VI—
VIII, in posterior band on segment V (para-
types sometimes with 1 or 2 pores near an-
terior margin of segment V and near pos-
terior margin of segment IV), without pores
on thorax (1 of 10 paratypes with | pore on
prothorax near anterior leg). Trilocular
pores scattered over surface. Discoidal
pores of same size as on dorsum, with 1
discoidal pore near eye on | side of body,
absent near other eye (paratypes with 1(0-—
2) pores near each eye. Oral-collar tubular
ducts of 1 size, present near marginal and
submarginal areas of abdomen and near se-
tal bases in medial and submedial areas of
abdomen and thorax, | oral collar mesad of
cerarius 12 (paratypes 2(0—5) ducts), with-
out oral collars in marginal or submarginal
areas of thorax or head. Setae as follows: 6
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
cisanal, (paratypes 4(3—6), longest 42 wp
long (paratypes 44(37—49) 2); longest anal-
lobe seta 148 w long (paratypes 157(124-
168) w); longest seta on trochanter 111 p
long (paratypes 109(99—124) w).
Circulus 96 w wide (paratypes 90(74-—
104) w), divided by intersegmental fold. La-
bium 161 yw long (paratypes 170(161—180)
w.). Antennae 8-segmented, (paratypes rare-
ly 7-segmented) 353 wp long (paratypes
341(316—366) w). Legs with 33 translucent
pores on hind femur (paratypes 29(15—52)
pores); 21 pores on hind tibia (paratypes
24(8—37) pores). Femur 212 w long (para-
types 210(185—235) w); tibia 198 pw long
(paratypes 190(170—217) w); tarsus 91 pw
long (paratypes 99(91—101) w). Tibia/tarsus
2.2 (paratypes 2.0(1.8—2.3)); femur/tibia 1.1
(paratypes 1.1(1.1—1.2)). Hind tibia with 19
setae (paratypes 19(16—23) setae). Length
of hind femur divided by greatest width of
femur 3.1(2.9-3.4). Claw digitules on hind
2 pairs of legs clubbed, each claw with 1
digitule with club slightly larger than club
on other digitule; claw digitules of front
pair of legs clubbed, about equal in size.
Tarsal digitules on hind 2 pairs of legs api-
cally clubbed, each tarsus with 1 digitule
with club noticeably larger than club on
other digitule; tarsal digitules on front pair
of legs of 2 different sizes and shapes, 1
digitule on each tarsus clubbed and nearly
reaching tip of claw, other digitule short
and apically acute.
Notes.—The above description is based
on 733 specimens from 4 localities. Adult
females can be distinguished from all other
instars by having multilocular pores, trans-
lucent pores on the hind femur and tibia,
and a vulva.
THIRD-INSTAR FEMALE
(Fig. 2)
Slide-mounted characters.—Body oval,
1.2(1.0-1.4) mm long, 0.8(0.6—0.9) mm
wide.
Dorsum with 17 pairs of cerarii, cerarii |
and 2 with 2 conical setae, cerarii 3 and 4
with 2(1—2) conical setae, cerarius 5 with 2
443
VOLUME 99, NUMBER 3
, Variety Farms, Atlantic County, New Jersey, XI-17-
Adult female Dysmicoccus vaccinii. Hammonton
Fig. 1.
1994, on Vaccinium corymbosum, S. Polavarapu.
tit
conical setae, cerarius 6 with 2(0—2) conical
setae, cerarius 7 with 2 conical setae, cer-
arius 8 with 2(0—2) conical setae, cerarius
9 with 2(1—2) conical setae, cerarius 10
with 2(0—3) conical setae, cerarius 11 with
2(1—2) conical setae, cerarius 12 with 3(2—
3) conical setae, cerarius 13 with 2(1-—2)
conical setae, cerarius 14 with 2(1—3) con-
ical setae, cerarius 15 with 3(2—3) conical
setae, cerarius 16 with 4(3—4) conical setae,
cerarius 17 with 3(3—4) conical setae. Cer-
arius 12 with 1(0—3) auxiliary setae, 10(7—
14) trilocular pores, and 1(1—2) discoidal
pores. Multilocular pores absent; trilocular
pores evenly scattered over surface; discoi-
dal pores about equal to diameter of triloc-
ular pore. Oral-collar tubular ducts absent.
Longest submedial seta on segment VII
24(19-28) ww long; 3(3—4) submedial setae
on segment VIII, longest seta 25(20—28) w
long.
Anal ring seta 93(86—-101) wp long;
1.5(1.4—1.7) times as long as greatest di-
ameter of ring.
Venter without multilocular pores. Tri-
locular pores scattered over surface. Dis-
coidal pores of same size as on dorsum,
with 1(0—2) pores near each eye. Normally
with | oral-collar tubular duct in cluster of
setae posterior of each spiracle; oral collars
absent elsewhere. Setae as follows: 4 cis-
anal setae, longest 31(25—37) w long; lon-
gest anal-lobe seta 125(111—138) w long;
longest seta on trochanter 71(54-82) w
long.
Circulus 62(49—74) ww wide, divided by
intersegmental fold. Labium 131(122—136)
w long. Antennae 6- or 7-segmented,
244(230—259) ww long. Legs without trans-
lucent pores. Femur 136(131—143) wp long;
tibia 105(96—109) yw long; tarsus 89(84—95)
w long. Tibia/tarsus 1.2(1.1—1.2); femur/tib-
ia 1.3(1.2—-1.4). Hind tibia with 10(8—11)
setae. Length of hind femur divided by
greatest width of femur 2.4(2.3—2.7). Claw
and tarsal digitules same as in adult female.
Notes.—This description is based on 313
specimens from 3 localities. The third-instar
female can be distiguished from all other
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
instars by having each cerarius with conical
setae, antennae usually 7-segmented, rarely
6, hind tibia length divided by hind tarsus
length 1.1 to 1.2, usually 1.2, and cerarius
12 with 7—14 associated trilocular pores. It
is most similar to the second-instar female
which differs by having cerarii anterior of
cerarius 7 without conical setae (cerarian
setae are filamentous), antennae usually
6-segmented, hind tibia length divided by
hind tarsus length 0.9, and cerarius 12 with
2-5 associated trilocular pores.
SECOND-INSTAR FEMALE
(Fig. 3)
Slide-mounted characters.—Oval, 0.9(0.8—
1.0) mm long, 0.5(0.5—0.6) mm wide.
Dorsum with 17 pairs of cerarii, posterior
cerarli to cerarius 7 or 8 usually with at
least 1 conical seta and | filamentous seta,
cerarli 1-6 with 2 setae, cerarii 7—9 with
2(1—2) setae, cerarius 10 with 2(1-—3) setae,
cerarius 11 with 2(1—2) setae, cerarius 12
with 3(2—3) setae, cerarius 13 with 2(1—2)
setae, cerarius 14 with 2(1—2) setae, cerar-
ius 15 with 3(2—3) setae, cerarius 16 with
2(1—2) setae, cerarius 17 with 3(3—4) setae.
Cerarius 12 with 3(2—5) trilocular pores,
and 1(0—1) discoidal pores. Multilocular
pores absent; trilocular pores scattered over
surface; discoidal pores about equal to di-
ameter of trilocular pores. Oral-collar-tu-
bular ducts absent. Longest submedial seta
on segment VII 17(15—22) w long; 1(0—2)
submedial setae on segment VIII, longest
seta 14(12—16) p long.
Anal ring seta 66(57-79) w long;
1.5(1.3—1.7) times as long as greatest di-
ameter of ring.
Venter without multilocular pores. Tri-
locular pores scattered over surface. Dis-
coidal pores of same size as on dorsum,
with 1(0—2) pores near each eye. Without
oral-collar tubular ducts. Setae as follows:
4 cisanal setae, longest 23(17—27) w long;
longest anal-lobe seta 93(84—99) w long;
longest seta on trochanter 71(54—-82) w
long.
Circulus 44(35—52) ww wide, divided by
VOLUME 99, NUMBER 3 445
Fig. 2. Third-instar female Dysmicoccus vaccinii. Hammonton, Variety Farms, Atlantic County, New Jersey,
XI-1-1995, on Vaccinium corymbosum, S. Polavarapu and D. R. Miller.
446 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
/
Fig. 3. Second-instar female Dysmicoccus vaccinii. Hammonton, Variety Farms, Atlantic County, New Jer-
sey, XI-1-1995, on Vaccinium corymbosum, S. Polavarapu and D. R. Miller.
VOLUME 99, NUMBER 3
intersegmental fold. Labium 99(94—104) w
long. Antennae 6-segmented, 244(230—
259) pw long. Legs without translucent
pores. Femur 91(86—99) w long; tibia
67(62—73) w long; tarsus 74(69—79) w long.
Tibia/tarsus 0.9; femur/tibia 1.4(1.3—1.4).
Hind tibia with 9 setae. Length of hind fe-
mur divided by greatest width of femur
2.2(2.0—2.4). Claw digitules same as on
adult female. Tarsal digitules on hind 2
pairs of legs with 1 digitule with club and
other apically acute and slightly shorter;
tarsal digitules on front pair of legs of about
same as other legs except apically acute
digitule is very short.
Notes.—This description is based on 78
specimens from 3 localities. The second-in-
star female can be distiguished from all oth-
er instars by having antennae usually 6-seg-
mented, hind tibia length divided by hind
tarsus length 0.9, and cerarius 12 with 2-5
associated trilocular pores. It is most similar
to the second-instar male which differs by
having oral-collar tubular ducts; these are
absent on the second-instar female.
FIRST INSTAR
(Fig. 4)
Slide-mounted characters.—Oval, 0.6(0.5—
0.7) mm long, 0.3(0.3—0.4) mm wide.
Dorsum with 16 pairs of definite cerarii,
anterior cerarius indefinite, posterior cerarii
to cerarius 2, 3, or 4 with at least 1 conical
seta and | filamentous seta, cerarii 1—9 with
2 setae, cerarius 10—16 with 2(1—2) setae,
cerarius 17 indefinite, represented by 1 or
more unassociated setae. Cerarius 12 with
1 trilocular pore. Multilocular and discoidal
pores absent; trilocular pores arranged in 4
longitudinal lines on each side of body.
Oral-collar tubular ducts absent. Longest
submedial seta on segment VII 10(8—11) wp
long; without submedial setae on segment
Vill.
Anal ring seta 52(47-54) w long;
1.7(1.6-1.9) times as long as greatest di-
ameter of ring.
Venter without multilocular pores. Tri-
locular pores arranged in 1 mediolateral
447
longitudinal line on each side of abdomen,
more abundant on thorax and head. Discoi-
dal pores associated with base of sublateral
line of setae, also with 1 associated with
each spiracle, with 1(0—1) pore near each
eye. Without oral-collar tubular ducts. Setae
as follows: 4 cisanal setae, longest 18(16—
22) pw long; longest anal-lobe seta 50(42—
61) pw long; longest seta on trochanter
39(32—44) wp long.
Inner circle of circulus 30(27-35) wp
wide, divided by intersegmental fold. La-
bium 76(64—83) w long. Antennae 6-seg-
mented, 142(128—-158) pw long. Legs with-
out translucent pores. Femur 63(59-—68) wp
long; tibia 46(42—49) pw long; tarsus 60(56—
65) w long. Tibia/tarsus 0.8(0.7—0.8);
femur/tibia 1.4(1.3—1.5). Hind tibia with 9
setae. Length of hind femur divided by
greatest width of femur 2.1(1.9—2.3). Tarsal
digitules on hind 2 pairs of legs with | dig-
itule with club and other apically acute and
slightly shorter; tarsal digitules on front pair
of legs of about same as other legs except
apically acute digitule is very short.
Notes.—This description is based on 70
specimens from 3 localities. The first instar
can be distiguished from all other instars by
having 6-segmented antennae, hind tibia
length divided by hind tarsus length 0.7—
0.8, usually 0.8, and cerarius 12 with 1 as-
sociated trilocular pore. It is most similar to
the second-instar female which differs by
having hind tibia length divided by hind
tarsus length 0.9, and cerarius 12 with 2—5
associated trilocular pores.
SECOND-INSTAR MALE
(Fig. 5)
Slide-mounted characters.—Body oval,
0.9(0.8-1.0) mm long, 0.6(0.5—0.6) mm
wide.
Dorsum with 17 pairs of cerarii, posterior
cerarii to cerarius 4 or 5 usually with at
least 1 conical seta and 1 filamentous seta,
cerarii 1—7 with 2 setae, cerarii 8 and 9 with
2(1-2) setae, cerarius 10 with 2(1—2) setae,
cerarius 11 with 2 setae, cerarius 12 with
2(1-3) setae, cerarius 13 with 2(1—2) setae,
448 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. First-instar (sex undetermined) Dysmicoccus vaccinii. Hammonton, Variety Farms, Atlantic County,
New Jersey, XI-1-1995, on Vaccinium corymbosum, S. Polavarapu and D. R. Miller.
cerarius 14 with 2(1—2) setae, cerarius 15 O(O—1) discoidal pores. Multilocular pores
with 2(1—3) setae, cerarius 16 with 2(2—3) absent; trilocular pores scattered over sur-
setae, cerarius 17 with 3(2—3) setae. Cer- face; discoidal pores about equal to diam-
arius 12 with 2(1-3) trilocular pores, and eter of trilocular pores. Oral-collar tubular
VOLUME 99, NUMBER 3
ducts of 1 size, same as large size on venter,
present in rows across segments beginning
on segment V or VI forward to head. Lon-
gest submedial seta on segment VII 16(12—
19) w long; 1(1—2) submedial setae on seg-
ment VIII, longest seta 14(10—20) wp long.
Anal ring seta 69(57-77) wp long;
1.5(1.3—1.7) times as long as greatest di-
ameter of ring.
Venter without multilocular pores. Triloc-
ular pores scattered over surface. Discoidal
pores of same size as on dorsum, with 1(0-—
1) pores near each eye. With 2 sizes of oral-
collar tubular ducts, larger size present in
margin or submargin from segments VII or
VI forward to head; smaller size in rows on
segments VII and VI and occasionally on
segment V, also present in medial and sub-
medial areas of anterior abdominal segments,
thorax, and head. Setae as follows: 4 cisanal
setae, longest 23(19—32) pw long; longest anal-
lobe seta 92(86—106) w long; longest seta on
trochanter 53(42—59) w long.
Circulus 38(35—42) w wide, divided by
intersegmental fold. Labium 95(91—99) w
long. Antennae 6-segmented, 177(153-—191)
w long. Legs without translucent pores. Fe-
mur 87(79—94) w long; tibia 66(56—69) w
long; tarsus 67(63—69) w long. Tibia/tarsus
1.0(0.9—1.0); femur/tibia 1.3(1.3—1.4). Hind
tibia with 9 setae. Length of hind femur di-
vided by greatest width of femur 2.3(2.1—
2.4). Claw digitules same as on adult fe-
male. Tarsal digitules on hind 2 pairs of
legs with | digitule with club and other api-
cally acute and slightly shorter; tarsal digi-
tules on front pair of legs of about same
dimensions as other legs except apically
acute digitule is very short.
Notes.—This description is based on 109
specimens from 3 localities. The second-in-
star male can be distiguished from all other
instars by having dorsal oral-collar tubular
ducts, mouthparts, and no vulva.
THIRD-INSTAR MALE (PREPUPA)
(Fig. 6)
Slide-mounted characters.—Body elon-
gate, 0.9 mm long, 0.4 mm wide.
449
Dorsum without cerarii, posterolateral mar-
gins of segments VI, VII, and VIII each with
2 setae conspicuously longer than remaining
setae on segments. Multilocular pores scat-
tered over surface except on mesothorax and
abdominal segments VII and IX; trilocular
pores absent; discoidal pores associated with
multilocular pores and oral collars. Oral-col-
lar tubular ducts of 1 size, scattered over sur-
face except on mesothorax and abdominal
segments VIII and IX. Longest submedial
seta on segment VII 17 p» long; 3 submedial
setae on segment VIII, longest seta 15 long.
Anal ring without setae and pores.
Venter with multilocular pores scattered
over surface except segments VIII and IX.
Trilocular pores absent. Discoidal pores as-
sociated with multiloculars and oral collars.
With oral-collar tubular ducts in marginal
areas except on abdominal segments VIII
and IX.
Circulus 82 wide, divided by interseg-
mental fold. Mouth structure weakly indi-
cated.Antennal segments indistinct, 232
long. Legs without translucent pores. Femur
91p long; division between tibia and tarsus
indistinct, tibiat+tarsus 148y long. Wing
buds of mesothoracic wings protruding
from lateral margin, about 110 long. Wing
buds of hamulohalterae absent.
Notes.—The above description is based
on | specimen reared in the Laboratory that
originally was collected in Hammonton,
New Jersey on Vaccinium corymbosum,
March 19, 1996. Preserved June 6, 1996. A
second specimen, field collected from the
same locality and host on September 28,
1996, lacks all signs of wing buds and has
a very weak indication of the circulus. We
suspect that this specimen is a prepupa of
the apterous male. The prepupa can be dis-
tiguished from all other instars by having,
multilocular pores, oral-collar tubular ducts,
antennae without definite segmentation, tib-
ia and tarsus fused, no mouthparts, no ae-
deagus, no definite constriction for the
head. It is most similar to the pupa which
differs by having antennae 10-segmented
and a definite constriction for head.
450 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 5. Second-instar male Dysmicoccus vaccinii. Hammonton, Variety Farms, Atlantic County, New Jersey,
XL-1-1995, on Vaccinium corymbosum, S. Polavarapu and D. R. Miller.
VOLUME 99, NUMBER 3 451
Fig. 6. Third-instar male (prepupa) Dysmicoccus vaccinii. Hammonton, Variety Farms, Atlantic County, New
Jersey, II-19-1996, on Vaccinium corymbosum, S. Polavarapu and D. R. Miller. Reared in laboratory VI-6-
1996.
FOURTH-INSTAR MALE (PUPA) on posterolateral margins of segments VI,
(Fig. 7) VII, and VIII each with 2 setae conspicu-
Slide-mounted characters—Body elon- ously longer than remaining setae on se-
gate, 1.0(0.9-1.1)mm long, 0.4(0.3— gements. Multilocular pores present in me-
0.4)mm wide.Dorsum without cerarii, setae diolateral areas of head, thorax, and abdo-
452
Fig. 7.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fourth-instar male (pupa) Dysmicoccus vaccinii. Hammonton, Variety Farms, Atlantic County, New
Jersey, II-19-1996, on Vaccinium corymbosum, S. Polavarapu and D. R. Miller. Reared in laboratory VI-6-
1996.
men, most abundant on prothorax; trilocular
pores absent; discoidal pores associated
with multilocular pores. Oral-collar tubular
ducts of | size, present in submarginal areas
of prothorax and abdomen. Longest sub-
medial seta on segment VII 29(25-30) pw
long; 5(4—5) submedial setae on segment
VIII, longest seta 30(25—32) pw long.
Anal ring without setae and pores.
Venter with multilocular pores in medio-
lateral areas of thorax and abdomen. Triloc-
ular pores absent. Discoidal pores associ-
ated with multiloculars. With oral-collar tu-
bular ducts in marginal areas of prothorax
and abdominal segments II or III to VII.
Postocular ridge and mesosternal furca ev-
ident.
Antennae 10-segmented, 368(353—384)
w long. Legs without translucent pores. Fe-
mur 117(111—121) p long; tibia 122(116—
128) pw long; tarsus 96(94—99) w long. Tib-
ia/tarsus 1.3(1.2—1.3); femur/tibia 0.9(0.9—
VOLUME 99, NUMBER 3
1.0). Wing buds of mesothorax protruding
from lateral margin, 289(248—347) w long.
Wing buds of hamulohalterae represented
by small protrusions on lateral margin of
metathorax.
Notes.—The above description is based
on 3 specimens reared in the laboratory that
originally were collected in Hammonton,
New Jersey, March 19, 1996 and preserved
June 6, 1996. We suspect that these speci-
mens are pupae of the macropterous form.
Specimens have also been collected near
Frankfort, Sussex Co., Delaware, April 5,
1996 (1); Hammonton, New Jersey, August
8, 1996 (1), September 28, 1996 (4), Oc-
tober 30, 1996 (1). In most of these speci-
mens, the apterous adult is inside. The pupa
does not have wing buds except for a slight-
ly wrinkled area where the mesothoracic
wing bud would be on the macropterous
form. The pupa can be distinguished from
all other instars by having multilocular
pores, oral-collar tubular ducts, 10-seg-
mented antennae, no mouthparts, no aede-
agus, definite constriction for the head. It is
most similar to the prepupa. For a compar-
ison see the notes section of the prepupa.
MACROPTEROUS ADULT MALE
(Fig. 8)
Slide-mounted characters.—Body elon-
gate oval, 1.0 mm long, 0.3 mm wide.
Dorsum with 1 pair of tail-forming pore
clusters; each cluster with 2 elongate setae
about 355y long, 1 or 2 additional shorter
setae, 32(30—34) multilocular pores, and 2
or 3 discoidal pores. Multilocular pores in
marginal areas of thorax and abdomen, with
4 or 5 loculi, quadriloculars most abundant.
Discoidal pores associated with multilocu-
lars, with 1 or 2 near base of antenna, oc-
casionally with | or 2 such pores in medial
areas of abdomen. Body setae of 2 kinds,
fleshy setae and bristle shaped; both kinds
scattered over surface. Abdominal scleroti-
zation restricted to abdominal tergite VIII.
Metapostnotal ridge conspicuous. Scutel-
lum rectangular, without scutellar ridge,
with several setae laterally. Scutum sclero-
453
tized throughout except with a median lon-
gitudinal clear area, area lateral of prescu-
tum heavily sclerotized, with reticulate pat-
tern, scutum with many setae. Prescutum
rectangular, with weakly defined prescutal
suture, with several setae along lateral and
posterior margins. Pronotal ridges heavily
sclerotized. Hamulohalterae 65(64—67) w
long, with 1 apical hooked seta. Mesotho-
racic wings 796(790-—883) w long, each
with 2 or 3 basal setae and 2 discoidal
pores. Dorsal arm of midcranial ridge ex-
tending to posterior margin of dorsal eye.
Dorsal medialsclerite sclerotized with nu-
merous setae. Dorsal eye about 32 in di-
ameter. Lateral ocellus 21(20—22) pw in di-
ameter, located at junction of preocular and
postocular ridges. Ocular sclerite lightly
sclerotized.
Penial sheath 147(146—-148) w long,
78(77—-79) w wide; length/width ratio 1.9.
Aedeagus 116(111—121) wp long, apically
truncate.
Venter with setae of same 2 shapes as on
dorsum, present medially, submedially and
laterally on most segments, abundant on
basisternum. Abdominal sclerotization con-
fined to sternite VIII. Prosternal ridge well
developed, sternite weakly sclerotized.
Preoral ridge weakly developed. Ocular
sternite sclerotized near ventral eye. Ventral
midcranial ridge well developed, with lat-
eral arms. Ventral eye about 37 in diam-
eter.
Hind femur 171(168—174) w long; tibia
211(210—212) w long; hind tarsus 90(86—
94) pw long; femur/tibia 0.8; tibia/tarsus
2.4(2.3—2.4). Slender fleshy setae present
on legs and antennae; apical segment of an-
tenna with capitate setae. Tarsal digitules
capitate; claw digitules acute. Antennae
10-segmented, 536(521—546) w long; seg-
ment 3 longest, 73(72—74) w long; segment
10, 70(69-72) w long; segment 3/10
1.0(1.0—1.1).
Notes.—The above description is based
on 2 specimens reared in the Laboratory
that originally were collected in Hammon-
ton, New Jersey on Vaccinium corymbos-
454 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Ne ny fl AR / A}
“7 rt; bik y
Ap Diss A
°
Fig. 8. Fifth-instar male (adult, macropterous form) Dysmicoccus vaccinii. Hammonton, Variety Farms, At-
lantic County, New Jersey, II-19-1996, on Vaccinium corymbosum, 8. Polavarapu and D. R. Miller. Reared in
laboratory VI-6-1996.
VOLUME 99, NUMBER 3
um, March 19, 1996 and preserved June 6,
1996. Another specimen with brachypter-
ous wings was reared at the same time. It
is virtually identical with the macropterous
forms but has small round wing buds and
no hamulohalterae. It is unclear if this is a
specimen that was in the process of wing
elongation or is a naturally occurring bra-
chypterous form. This macropterous form
of the adult male can be distinguished from
all other instars by possessing wings, a def-
inite aedeagus, lateral pore clusters, and a
heavily sclerotized thorax and head.
APTEROUS ADULT MALE
(Fig. 9)
Slide-mounted characters.—Body elon-
gate oval, 1.2(1.0—1.2) mm long, 0.4 mm
wide.
Dorsum with 1 pair of tail-forming pore
clusters; each cluster with 2 elongate setae
276(260—291) w long, 1 or 2 additional
shorter setae, 31(25—37) multilocular pores,
and 2 or 3 discoidal pores. Multilocular
pores in marginal areas of head, thorax and
abdomen, with 3, 4, or 5 loculi, quadrilo-
culars most abundant. Discoidal pores as-
sociated with multiloculars, with several
near base of antenna, occasionally with | or
2 such pores in medial areas of abdomen.
Body setae of 2 kinds, fleshy setae and bris-
tle shaped; fleshy setae on abdomen, bristle-
shaped setae scattered over surface. Ab-
dominal sclerotization most conspicuous on
abdominal tergite VIII and in lateral areas,
weakly evident in mendial and submedial
areas. Metapostnotal ridge inconspicuous.
Scutellum, scutum, and prescutum fused
into | sclerotized area. Hamulohalterae and
mesothoracic wings absent. Dorsal arm of
midcranial ridge variable, extending to pos-
terior margin of dorsal eye in some speci-
mens, represented by weak sclerotiztion on
others. Dorsal medialsclerite unsclerotized
with numerous setae, discoidals, and a few
multilocular pores. Dorsal eye about 25(22—
27) w in diameter. Lateral ocellus 24(22—
27) w in diameter, located at junction of
455
preocular and postocular ridges. Ocular
sclerite lightly sclerotized.
Penial sheath 156(148-161) w long,
87(79-91) ww wide; length/width ratio
1.8(1.7—2.0). Aedeagus 121(111-131) p
long, apically truncate.
Venter with setae of same 2 shapes as on
dorsum, present medially, submedially and
laterally on most segments, abundant on
basisternum. Abdominal sclerotization con-
fined to sternite VIII. Basisternum with an-
terior marginal ridge incomplete. Prosternal
ridge well developed, sternite weakly scler-
otized. Preoral ridge weakly developed. Oc-
ular sternite sclerotized near ventral eye.
Ventral midcranial ridge well developed,
with lateral arms. Ventral eye about 33(30—
35) w in diameter.
Hind femur 181(172—-191) wp long; tibia
210(200—221) ww long; hind tarsus 94(91—
99) w long; femur/tibia 0.8(0.8—0.9); tibia/
tarsus 2.2(2.1—2.3). Slender fleshy setae
present on legs and antennae; apical seg-
ment of antenna with capitate setae. Tarsal
digitules capitate; claw digitules acute. An-
tennae 9- or 10-segmented, when 9-seg-
mented, segments 4 and 5 fused, 486(477—
502) w long; segment 3 longest, 65(62—70)
ww long; segment 10(9), 62(59—65) w long;
segment 3/10(9) 1.1(1.0—1.1).
Notes.—The above description is based
on 5 specimens reared in the Laboratory
that originally were collected in Hammon-
ton, New Jersey on Vaccinium corymbos-
um, March 19, 1996 and preserved June 6,
1996. This form of the adult male can be
distinguished from all other instars by hav-
ing a definite aedeagus, lateral pore clusters,
and a heavily sclerotized thorax and head,
and by lacking wings.
SPECIMENS EXAMINED
Paratypes—DELAWARE: Near Frank-
ford, Sussex County, IV-5-1996, on Vac-
cinium spp., S. Polavarapu and D. R. Miller
(87 ad 2, 48 third-instar 2, 5 second-instar
?, 1 fourth-instar pupal ¢, 28 second-instar
3, 5 first instars) USNM. NEW JERSEY:
Near Hammonton, Variety Farms, Atlantic
456 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
. oy.
STE SIS/ Wht
fT Fil ye
ve i NIT ii Yee
Fig. 9. Fifth-instar male (adult, apterous form) Dysmicoccus vaccinii. Hammonton, Variety Farms, Atlantic
County, New Jersey, III-19-1996, on Vaccinium corymbosum, S. Polavarapu and D. R. Miller. Preserved after
rearing in laboratory VI-6-1996.
County, V-12-1994, VI-7-1994, XI-17- IV-5-1996, IV-23-1996, V-6-1996, Vaccin-
1994, XII-30-1994, I-13-1995, III-14-1995, ium spp., (XI-1-1995 infestation also found
IV-14-1995, XI-1-1995, I-13-1995, II-12- on Polygonum sp.) D. R. Miller and/or S.
1996, III-5-1996, III-19-1996, III-20-1996, Polavarapu ( 560 ad @, 235 third-instar °,
VOLUME 99, NUMBER 3
73 second-instar °, 2 macropterous ad d,
1 brachypterous ad 6, 5 apterous ad d, 3
fourth-instar pupal 6, 1 third-instar prepu-
pal d, 75 second-instar d, 64 first instars)
BMNH, CDA, FSCA, MHNH, UCD,
USNM;; Near Hammonton, MacCrie Broth-
ers Farm, Atlantic County, III-5-1996, Vac-
cinium spp., S. Polavarapu and D. R. Miller
(100 ad 2, 30 third-instar 2, 6 second-in-
star 6, 1 first instar) USNM; Near Ham-
monton, MacCrie Brothers Farm, Atlantic
County, VII-22-1993, IX-7-1993, [X-29-
1993, Vaccinium spp., K. S. Samoil (24 ad
2?) USNM.
Not paratypes—NORTH CAROLINA:
Bailey, Nash County, I-25-1973, on Vaccin-
ium ashei, H. H. Neunzig (6 ad 6) USNM.
There is an additional series of speci-
mens from the Hammonton locality that
were collected August 8, 1996, September
28, 1996, and October 30, 1996. This ma-
terial was mounted quickly for assessing
the presence of different stages of the
mealybug and therefore is not included in
the type series. It includes 396 adult fe-
males, 139 third-instar females, 31 second-
instar females, 111 first instars, 1 apterous
adult male, 7 pupal fouth-instar males, |
prepupal third-instar male, and 35 second-
instar males. All of this material is depos-
ited in the USNM.
Specimens collected in North Carolina
are believed to be conspecific with Dysmi-
coccus vaccinii but have shorter append-
ages and shorter dorsal setae and therefore
are not included in the type series. The sub-
mittal slip from H. H. Neunzig, North Car-
olina State reads “‘I am sending specimens
collected from rabbit-eye blueberries (Vac-
cinium ashei) at a nursery in Bailey, N. C.
They occur in large numbers and are form-
ing galls on the roots. A white secretion is
also associated with these insects.’’ In New
Jersey infestations we have never seen any
indication of galls and have been unable to
confirm their existence in North Carolina.
An additional series of specimens includ-
ing 9 adult females, 3 third-instar females,
and 4 first instars, was submitted to the Sys-
457
tematic Entomology Laboratory in 1984 by
Donald Ramsdel!l of Michigan State Uni-
versity. Data on the slides indicate only that
the specimens were collected on blueberry
in New Jersey in November 1984. Since we
do not have specific locality information,
this series has not been included in the type
series. The specimens fall well within the
range of variation of Dysmicoccus vaccinii.
Field results—Sampling of field popu-
lations during late fall of 1995 through fall
of 1996 revealed the presence of various
immature and adult stages throughout the
sampling period (Fig. 10). Although sec-
ond-instar males were consistently present,
the adult males were found in the field only
four times. One pupa was collected near
Frankford, DE on April 5, 1996. In Ham-
monton, NJ male stages after the second in-
star were collected in the field on the fol-
lowing dates: one pupa (August 8, 1996);
one prepupa, 4 pupae, and one apterous
adult male (September 28, 1996); 2 pupae
(October 30, 1996).
Adult females collected from the field
during January and February readily ovi-
posit in the laboratory. Eggs are laid within
an ovisac made of wax filaments. Dysmi-
coccus vaccinii apparently has more than
one generation each year in New Jersey.
In our field observations, we have always
found D. vaccinii populations associated
with the ants, Acanthomyops claviger (Rog-
er) or Lasius neoniger Emery. These ant
species were seen tending all stages of
mealybugs. Ants were also seen carrying
mealybugs especially in response to distur-
bance.
DISCUSSION AND CONCLUSIONS
Dysmicoccus vaccinii is part of a com-
plex of mealybugs characterized by Beards-
ley (1965) as the D. brevipes group or the
pineapple mealybug complex. Morpholog-
ically, they are recognized as species of
Dysmicoccus that have most of the follow-
ing characters: discoidal pores near the rim
of the eye, multilocular pores restricted to
the ventral surface of the abdomen, a cir-
458
80
@ First-instar
if S Second-instar (male)
= ® Second-instar (female)
os ® Third-instar (female)
a 60 % Fourth-instar (male)
~ Z Adult (male)
8 S Adult (female) \
:
3 \
« 40 \ \
S \ N
E \
=
a \
= \
- \ & XN
~ 20 5
5 aN
2 & N
3 N & NV
“f \ \ | \
_\
aN
; Ne
1-Nov.95 12-Feb.96 5-Mar.96
5-Apr.96
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
per ay
\
\ \ \ \
N N N
\ \ \
N S
N N N N
N N N N \N
\N N N N N
Wittman
Bilal
\ \
Biiminnne
N
GY: aN \
5 N
\y% \ ae, aN
23-Apr.96 8-Aug.96 28-Sep.96 30-Oct.96
Dates
Fig. 10. Distribution of immature and adult stages of D. vaccinii on blueberries during November, 1995 to
October, 1996, near Hammonton, New Jersey. Third-instar males are not included but one specimen was collected
9/28/96 and is 0.8% of total for that collection. Sample size was 65—439 mealybugs on different sampling dates.
culus divided by the intersegmental line, 17
pairs of cerarii, and translucent pores on at
least the hind femur and tibia. With the ad-
dition of Dysmicoccus vaccinii to the group
there now are 12 species in the complex
including D. brevipes, D. grassii (Leonar-
di), D. mackenziei Beardsley, D. morrisoni
(Hollinger), D. neobrevipes Beardsley, D.
pinicolus McKenzie, D. probrevipes (Mor-
rison), D. radicis (Green), D. roseotinctus
(Cockerell and Cockerell), D. texensis, and
D. tibouchinae (Hambleton).
The adult female of Dysmicoccus vacci-
nii is very similar to D. texensis, but differs
by having no oral collars on the head, 1 size
of oral collar on the body, the length of the
hind femur divided by greatest width of the
femur 3.1(2.9—3.4), and the longest dorso-
medial seta on segment VIII 34(25—42) wp
long. Dysmicoccus texensis, has several oral
collars near the anterior margin of the head,
2 distinct sizes of oral collars, the length of
hind femur divided by the greatest width of
femur 2.0(1.9—2.1), and the longest dorso-
medial seta on segment VIII 21(20—23) wp
long.
The adult macropterous male is similar
in appearance to the adult male of Dysmi-
coccus grassii (=Dysimcoccus alazon Wil-
liams) as described by Afifi (1968) but has
an apically blunt aedeagus whereas D. gras-
sii has an apically acute aedeagus. It also is
similar to Dysmicoccus brevipes (Cocker-
ell) and D. neobrevipes Beardsley as de-
scribed by Beardsley (1965). Dysmicoccus
brevipes and D. neobrevipes differ by hav-
ing the apex of the aedeagus bifurcate; D.
neobrevipes also has numerous short, en-
VOLUME 99, NUMBER 3
larged setae on the antennae which do not
occur in D. vaccinii, D. grassii, or D. brev-
ipes.
The key to adult females of North Amer-
ican species of mealybugs presented by
Miller and McKenzie (1973) needs to be
modified as follows to accommodate Dys-
micoccus vaccinil.
27(26). Oral-collar tubular ducts with dermal or-
ifices noticeably larger in diameter than
trilocular pores; lateral abdominal clusters
of oral-collar tubular ducts each with few-
er than 10 ducts .... aurantius (Cockerell)
- Oral-collar tubular ducts with dermal or-
ifices smaller than or equal to diameter of
trilocular pores; lateral abdominal clusters
of oral-collar tubular ducts each with
more than 10 ducts
28(27). Without oral-collar tubular ducts on head;
with only | size of oral collar
Beret On tai tee vaccinii Miller and Polavarapu
— With oral-collar tubular ducts on head;
With) ZISIZeSuOMmoralecOllarsies sn) ees oe
One morphological result merits further
discussion. As has been suggested previ-
ously (Miller 1975) the value derived from
dividing the hind tibia length by the hind
tarsus length is distinctive for each imma-
ture instar. In D. vaccinii these values are:
adult female 2.0(1.8—2.3); third-instar fe-
male 1.2(1.1—1.2); second-instar female
0.9; second-instar male 1.0(0.9—1.0); first-
instar 0.8(0.7—0.8). The interesting obser-
vation is that the value derived from divid-
ing the hind femur length by the hind tibia
length is basically the same for all instars.
In D. vaccinii these values are: adult female
1.1(1.1—-1.2); third-instar female 1.3(1.2-—
1.4); second-instar female 1.4(1.3—1.4);
second-instar male 1.3(1.3—1.4); first-instar
1.4(1.3—1.5); only the adult female differs
from the rest. Therefore, it appears that the
tarsus is growing at a different rate than the
other measurable segments of the leg.
In many insects, the diapausing stage is
species-specific, and is reached prior to the
arrival of adverse environmental conditions
(Tauber et al. 1986). The occurrence of im-
mature and adult stages of D. vaccinii
459
throughout the fall and winter suggests the
possibility that this insect does not have a
true overwintering stage, although this does
not preclude the possibility of overlapping
diapausing and non-diapausing generations.
For instance, the early-instar nymphs and
mature adult females found during late fall
and winter may represent diapausing and
non-diapausing generations, respectively.
Mature females collected during winter
readily resume oviposition in the laboratory
at 20-23°C. This suggests that mature
mealybugs are in a state of quiescence
awaiting the onset of favorable tempera-
tures.
In the present study, the majority of the
sampled population consisted of adult fe-
males at most times of the year. This may
be partially attributed to the sampling bias
against the immature stages of the mealy-
bug population. Nevertheless, field samples
collected throughout the study always con-
sisted of immature stages including second-
instar males. There may be a trend towards
an increasing percentage of third-instar and
adult females as winter progresses, but a
more rigorous sampling regime is required
to confirm this observation. Our failure to
collect adult males on a regular basis, in
spite of the common occurrence of second-
instar males in the field, is difficult to ex-
plain. It is entirely possible that we simply
are not locating the adults because of their
small size and short life span. Clearly, more
work is needed to understand the biology
and seasonal life-history of D. vaccinii.
Ant-mealybug mutualistic relationships
have been previously documented in the ge-
nus Dysmicoccus (e.g., Carter 1932; Milli-
ron 1958; Beardsley et al. 1982; Rohrbach
et al. 1988). Several species of ants feed on
honeydew produced by the gray pineapple
mealybug, Dysmicoccus neobrevipes. The
ants are reported to benefit this mealybug
species by providing protection from natu-
ral enemies and adverse weather conditions,
by transporting the mealybugs among
plants, and by removing honeydew which
prevents sooty mold buildup (Rohrbach et
460
al. 1988). Recent studies under laboratory
conditions, however, failed to demonstrate
the role of the big-headed ant, Pheidole
megacephala (F) in increasing the mealy-
bug colony size or in transporting the meal-
ybugs (Jahn and Beardsley 1996). Future
research should evaluate the role of ants in
maintaining populations of D. vaccinii in
blueberry fields, to determine whether
mealybug control strategies involving the
management of ant populations should be
developed and implemented.
ACKNOWLEDGMENTS
Thanks to Ms. Elizabeth Bender, Blue-
berry and Cranberry Research Center, Rut-
gers University (BCRC) for assisting in col-
lection of field samples. We also thank Dr.
Robin Stuart, BCRC for collecting some of
the male mealybug specimens and for his
comments and observations on ant-mealy-
bug interactions. We are grateful to David
R. Smith, Systematic Entomology Labora-
tory, Agricultural Research Service, United
States Department of Agriculture, Washing-
ton, D.C. (SEL), for identifying the ant spe-
cies. To Mr. John Bertino of Variety Farms
we gratefully acknowlege his willingness to
let us sample mealybugs in his blueberry
fields. This research was partially funded by
USDA-ARS-CSREES (93-34155-8382)
awarded to SP.
We are grateful to the following individ-
uals for reading and commenting on the
manuscript: Dr. John A. Davidson, Depart-
ment of Entomology, University of Mary-
land, College Park; Dr. William E Gimpel,
Jr., Plant Protection Section, Maryland De-
partment of Agriculture, Annapolis; Dr. Mi-
chael E. Schauff and Dr. E. Eric Grissell of
SEL. We also wish to acknowledge the im-
portant contribution of Ms. Debra Creel of
SEL who prepared more than a thousand
specimens of this new species.
LITERATURE CITED
Afifi, S. A. 1968. Morphology and taxonomy of the
adult males of the families Pseudococcidae and
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Eriococcidae. Bulletin of the British Museum
(Natural History) Entomology 13: 1—210.
Beardsley, J. W. 1965 (1964). Notes on the pineapple
mealybug complex, with descriptionsof two new
species. Proceedings of the Hawaiian Entomolog-
ical Society 19: 55-68.
Beardsley, J. W., T. H. Su, FE L. McEwen, and D. Ger-
ling. 1982. Fieldinvestigations on the interrela-
tionships of the big-headed ant, the gray pineap-
plemealybug, and pineapple mealybug wilt dis-
ease in Hawaii. Proceedings of the Hawaiian-
Entomological Society 24: 51—67.
Ben-Dov, Y. 1994. A systematic catalogue of the
mealybugs of the world. InterceptLimited, Ando-
ver, United Kingdom, 686 pp.
Carter, W. 1932. Studies of populations of Pseudo-
coccus brevipes (Ckll.) occurring onpineapple
plants. Ecology 13: 296-304.
Gimpel, W. EF, Jr. and Miller, D. R. 1996. Systematic
analysis of the mealybugs in the Pseudococcus mar-
itimus complex (Homoptera: Pseudococcidae). Con-
tributions on Entomology, International 2: 1-163.
Jahn, G. C. and J. W. Beardseley. 1996. Effects of
Pheidole megacephala (Hymenoptera: Formici-
dae) on survival and dispersal of Dysmicoccus
neobrevipes (Homoptera: Pseudococcidae). Jour-
nal of Economic Entomology 89: 1124-1129.
Miller, D. R. 1975. A revision of the genus Hetero-
coccus Ferris with a diagnosis of Brevennia Goux.
United States Department of Agriculture, Techni-
cal Bulletin Number 1497, 61 pp.
Miller, D. R. and McKenzie, H. L. 1973. Seventh tax-
onomic study of North Americanmealybugs. Hil-
gardia 41: 489-542.
Milliron, H. E. 1958. Economic importance and control
of the loblolly mealybug, Dysmicoccus obesus Lob.
Journal of Economic Entomology 51: 555-556.
Ramsdell, D. C., K. S. Kim, and J. P. Fulton. 1987.
Red ringspot of blueberry, pp 121—123. Jn Con-
verse, R. H., ed., Virus diseases of small fruits.
United States Department of Agriculture, Agri-
culture Handbook No 631. U. S. Government
Printing office, Washington, D.C., 277 pp.
Rohrbach, K. G., J. W. Beardsley, T. L. German, N. J.
Reimer, and W. G. Sanford. 1988. Mealybug wilt,
mealybugs, and ants on pineapple. Plant Disease
72: 558-565.
Tauber, M. J., C. J. Tauber, and S. Masaki. 1986. Sea-
sonal adaptations of insects. OxfordUniversity
Press, New York, New York. 411 pp.
Tinsley, J. D. 1900. Contributions to coccidology.—
II. Canadian Entomologist 32: 64—67.
Williams, D. J. and Granara de Willink, M. C. 1992.
Mealybugs of Central and South America. CAB In-
ternational, Wallingford, United Kingdom, 635 pp.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 461-471
REDESCRIPTION OF ANOPHELES (ANOPHELES) SHANNONI DAVIS;
A MEMBER OF THE ARRIBALZAGIA SERIES FROM THE
AMAZON BASIN (DIPTERA: CULICIDAE)
RICHARD C. WILKERSON, MARIA ANICE M. SALLUM, AND OSWALDO PAULO FORATTINI
(RCW) Department of Entomology, Walter Reed Army Institute of Research % Mu-
seum Support Center, Smithsonian Institution, Washington, D.C. 20560, U.S.A.; (MAMS,
OPF) Departamento de Epidemiologia, Faculdade de Saude Publica, NUPTEM, Univer-
sidade de Sao Paulo, Av. Dr. Arnaldo, 715, CEP 01246-904, Sao Paulo, S.P., Brazil.
Abstract.—Anopheles (Anopheles) shannoni Davis is redescribed and illustrated in the
adult female, male genitalia, and larval and pupal stages. This species is distributed
throughout the Amazon Basin of South America. The larvae are found in shaded forest
pools, and the adults appear to be zoophilic and prefer to bite outdoors. This species is a
member of the Neotropical Arribalzagia Series.
Key Words:
zon Basin
Anopheles (Anopheles) shannoni Davis is
a member of the Arribalzagia Series (Reid
and Knight 1961). Wilkerson and Peyton
(1990) implied that the approximately 23
species in this Neotropical group are mono-
phyletic based on shared wing spot char-
acters. Except for An. vestitipennis Dyar
and Knab (Belkin et al. 1970), An. male-
factor Dyar and Knab, An. punctimacula
Dyar and Knab (Wilkerson 1990) and An.
calderoni Wilkerson (Wilkerson 1991),
these species are not adequately described.
This redescription is part of an incremental
effort to characterize the species in the
group. In the following redescription Har-
bach and Knight (1980, 1982) were used
for morphological terminology and num-
bering of larval and pupal setae and, Wilk-
erson and Peyton (1990) for wing spot no-
menclature. Generic and subgeneric abbre-
viations follow Reinert (1975). An asterisk
in a taxonomic citation indicates illustration
of a given developmental stage. The spe-
cific feature(s) illustrated follows in brack-
ets.
Diptera, Culicidae, Arribalzagia, Anopheles shannoni, redescription, Ama-
Anopheles (Anopheles) shannoni Davis
Davis 1931: 345 (female* [wing, hind-
leg]). State of Para, Brazil. Holotype female
(National Museum of Natural History,
Washington, DC).
Female (Fig. 1).—Integument pale
brown to dark brown, grayish brown pol-
linose. Head: Interocular space with 6—9 (n
= 10 for this and following measurements
and counts except where indicated) long,
white setae and row of small, narrow, ap-
pressed pale yellow scales; vertex, occiput
and upper portion of postgena with numer-
ous erect, truncate scales; a patch of grayish
white to pale yellow scales on dorsal area
of vertex, nearly concolorous with scales on
median area of anterior promontory, a patch
of dark scales laterally on head concolorous
with scales on lateral area of anterior prom-
ontory and upper antepronotum; head with
15-23 long, black ocular setae; postgena
with long black setae ventrally. Clypeus
bare. Pedicel of antenna with 4—10 small,
dorsolateral, narrow to broad, grayish white
462
Pigade
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
CLES
Anopheles shannoni. Adult female habitus.
VOLUME 99, NUMBER 3
spatulate scales; flagellomere 1 with nu-
merous narrow to broad, dark spatulate
scales. Scales of maxillary palpus slender
and spatulate, mostly dark brown with in-
termixed dark brown setae; scales on pal-
pomeres 2—4 erect; pale yellow scales pres-
ent on the bases of palpomeres 3, 4 and 5;
length of maxillary palpus 1.99—2.61 mm
(mean 2.27 mm); ratio of length of palpom-
eres 2—5 to total length of palpus, 2 = 0.26—
0.35 (mean 0.31), 3 = 0.32—0.37 (mean
0.35), 4 = 0.17-0.22 (mean 0.20), 5 =
0.12—0.17 (mean 0.15); ratio of palpomere
4 to 5, 1.10—1.65 (mean 1.35); palpus 1.01—
1.25 (mean 1.12) forefemur length. Probos-
cis with dark brown setae and decumbent
dark brown scales, base with longer erect
scales and setae; proboscis length 2.21—2.81
mm (mean 2.50 mm), proboscis 1.03—1.16
(mean 1.10) palpus length. Thorax: Integ-
ument brown to dark brown, silvery polli-
nose. Scutum with 3 prominent dark brown
spots, 2 at the ends of and slightly posterior
to prescutal sutures and another in prescu-
tellar area continuing onto scutellum, some-
times 2 smaller spots at the ends of lateral
portions of prescutellar area. Scutal setae
numerous, pale yellow with golden reflec-
tions; scutum mottled with small dark
brown spots mostly corresponding to setal
insertions in acrostichal and dorsocentral
areas; median anterior promontory with
patch of long, narrow falcate white to pale
yellow scales; scutal fossa without scales
except anterior scutal fossa with patch of
broad, spatulate dark brown erect scales,
sometimes with a few intermixed pale
scales; supraalar area with spatulate, elon-
gate, narrow falcate, pale yellow scales.
Scutellum with 11—23 shorter and 14—20
long, pale yellow setae. Antepronotum with
20—43 yellowish to dark brown setae and
10-20 upper, dark spatulate scales. Pleural
vestiture as follows, with all scales white or
pale yellowish white and spatulate: upper
proepisternum with 3-6 setae, rarely with 1
narrow scale; prespiracular area with 6—13
setae, a single scale sometimes present;
prealar area with 10—24 setae; upper me-
463
sokatepisternum with 3-6 setae, rarely with
1 narrow scale; lower mesokatepisternum
with 2—5 setae, 7-9 scales; upper mesepi-
meron with 6—14 setae, 0—4 scales. Legs as
figured, scales dark brown and white or
pale yellow, scales and setae at apices of
fore- and hindtibiae yellow. Distribution of
scales on coxae and trochanters as figured.
Extent and number of pale spots on femora
and tibiae variable. Bases and apices of
femora pale; mid- and hindfemora with
ventral pale stripes, stripe on hindfemur dis-
tinct and with well demarcated borders;
fore- and midtibiae with ventral longitudi-
nal stripe of yellow scales, anterior, poste-
rior and dorsal surfaces with yellow spots,
hindtibia with an anterior longitudinal stripe
of yellow scales and an indistinct stripe of
pale yellow scales on posterior surface, ba-
sal portion with spots of yellow scales;
foretarsomere | with a ventral stripe of pale
scales, apex of foretarsomeres 1-5 with
pale spots, more evident on anterior sur-
face, 5 sometimes totally pale, midtarso-
mere | with indistinct pale stripe on ventral
surface, apex of tarsomeres 1—5 with pale
spots, these more evident anteriorly, ante-
rior surface of hindtarsomere | with an in-
distinct longitudinal stripe of pale scales
and with a few spots of pale scales at base,
hindtarsomere 2 with intermixed pale
scales, hindtarsomeres 1—5S with pale rings
at apices. Forefemur length 1.76—2.28 mm
(mean 2.03 mm), ratio of forefemur length
to proboscis length 1.16—1.29 (mean 1.23).
Wing (Table 1). Length (measured from hu-
meral crossvein) 3.47—4.52 mm (mean 3.93
mm). Dark scales brown to black, pale
scales nearly white. Basal pale spot usually
present; prehumeral pale spot absent; sub-
costa basad of humeral crossvein with patch
of dark scales ventrally, sometimes also
with a few white scales; humeral crossvein
dark-scaled dorsally and ventrally; acces-
sory sector dark spot present; often 2 pre-
and 2 postsubcostal pale spots and 1 pre-
and 1 postsubcostal dark spot, less often a
presubcostal and/or a postsubcostal pale
spot and/or a postsubcostal dark spot ab-
464
Table 1. Anopheles shannoni: descriptive statistics
for ratios of costal wing spot lengths to length of wing
measured from the humeral crossvein (n = 10 wings
from 10 individual females).
Wing Spot Range Mean SD
Basal pale 0.00-0.01 0.01 0.00
Prehumeral dark 0.08-0.11 0.10 0.01
Humeral pale 0.01-0.02 0.02 0.00
Humeral dark 0.04-0.07 0.06 0.01
Presector pale 0.01-0.02 0.02 0.00
Presector dark 0.07-0.12 0.10 0.01
Sector pale 0.07-0.13 0.09 0.02
Accessory sector dark 0.13-0.37 0.24 0.02
Sector dark 0.13-0.17. 0.15 0.02
Subcostal area 0.18-0.27 0.22 0.03
Presubcostal dark 0.00-0.06 0.03 0.02
Presubcostal pale (prox.) 0.01-0.04 0.02 0.01
Presubcostal pale (distal) 0.00-0.02 0.01 0.01
Postsubcostal dark 0.00-0.04 0.02 0.02
Postsubcostal pale (prox.) 0.04-0.05 0.02 0.02
Postsubcostal pale (distal) 0.00-0.03 0.01 0.01
Subcostal dark 0.08-0.12 0.10 0.01
Preapical dark 0.14-0.18 0.15 0.01
Preapical pale 0.01-0.05 0.03 0.01
Apical dark 0.00-0.10 0.02 0.04
sent; apical dark spot usually absent, occa-
sionally represented by a few scales at the
end of vein R, and sometimes dark scales
on fringe. Spots on posterior veins variable;
R,,; Sometimes mostly dark-scaled with in-
termixed pale scales, always with a small
pale spot and distinct black spot at proximal
end of vein, M, sometimes dark-scaled with
intermixed pale scales, sometimes with a
pale spot on proximal portion, sometimes
with 2 black spots, one on proximal and
another on distal end of vein, mcu at CuA
variable from white-scaled to dark-scaled.
Pale fringe spots indistinct, not well de-
marcated. Halter. Scabellum and ventral
surface of pedicel with pale integument,
dorsal surface of pedicel and capitellum
with brown integument; pedicel and capi-
tellum white-scaled dorsally, capitellum
dark-scaled ventrally, concave center with-
out scales. Abdomen: Integument brown to
dark brown with some grayish pollinosity.
Terga with numerous long yellowish setae;
terga II-VII with erect, posterolateral, dark
scale patches; tergum VIII with narrow,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
nearly white to yellow spatulate scales and
also with patches of posterolateral, dark
spatulate scales. Cercus distinctly constrict-
ed apically, pale yellow scaled. Sterna with
scattered brown to yellow setae; sternum I
without scales; sterna II—-VII with scattered
broad, white, spatulate scales and postero-
mesal patches of brown, dark, spatulate
scales; sternum VIII with scattered, narrow,
pale yellow and dark scales.
Male (Fig. 2).—As in female except for
the following sexual differences. Maxillary
palpus about 0.90 length of proboscis; apex
of palpomere 3 and all palpomeres 4 and 5
enlarged, palpomere 4 about 4 times broad-
er than base of palpomere 3. Maxillary pal-
pus with dark brown and white to pale yel-
low scales; basal 0.5 of palpomere 2 with
erect scales, apex with dorsolateral patch of
white scales; palpomere 3 dark-scaled with
an incomplete ring of white scales at base,
a dorsolateral patch of white scales on basal
0.3 and a few scattered pale yellow scales;
palpomere 4 mostly dark-scaled with scat-
tered, pale yellow scales on dorsal and lat-
eral surfaces and a patch of white scales at
apex; palpomere 5 mostly dark-scaled with
a dorsal patch of pale yellow scales at apex;
palpomeres 4 and 5 mostly bare mesally,
with long yellowish setae dorso- and ven-
tromesally, 5 with scattered pale yellow
scales. Proboscis length 2.87 mm, with
small, decumbent, dark brown scales, and a
ventrobasal patch of erect, dark scales, la-
bella brown. Foreungues with curved sub-
median tooth and short, blunt, external ba-
sal tooth. Genitalia: Ninth tergal lobes
short, somewhat triangular in outline, wide-
ly separated. Dorsal surface of gonocoxite
with a few scattered, moderately long setae,
lateral surface with slender fusiform and
spatulate scales, ventral surface as dorsal
surface but with lateral scales, most mesal
parabasal spine stout with slender, recurved
tip, borne on a slightly raised base; the oth-
er parabasal longer and more slender, both
about 0.23 from base of gonocoxite; inter-
nal seta slender, about as long as most mes-
al parabasal, base about 0.75 distance from
VOLUME 99, NUMBER 3 465
—1_.
Fig. 2. Anopheles shannoni. Pupa and male genitalia. GC—gonostylar claw, Gs—gonostylus, Gc—goncox-
ite, Cl—claspette, IX-Te—tergum IX, Ae—aedeagus, T—trumpet.
466
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 2. Pupal setal branching for Anopheles shannoni: range, mode (_). Most are based on counts of twenty
setae. In the case of equal modes, the highest is given. A range shown without a mode indicates that it was
indefinite. Seta 8—II was found in only one specimen.
Cephalo- Abdominal Segments
Seta thorax Paddle
No. Cr I Il Il IV Vv VI Vil Vill Ix
0) — — 14 (3) 1-7(3) 1-6(4) 1-5 (4) 2-6(4) 2-6(©4) 1 — a
] 2-4 (3) 13-23 (22) 5-10 (6) 3-7 (4) 2-5 (3) 1-3() 1-30) 1,2d) 1 1 PG)
2. 2376)) 2 0K@) 6-8 (7) 5-8 (6) 3-5 (4) 3-5 (4) 46(5) 3-6 (4) oo — 1-3 ()
3) 2-418) L2@® 2-5 (4) 3-7 (4) 410(8) 3-7 (6) 2-6 (3) 3-8 (6) — — —
4 2-5 (4) 2-8 (4) 2-6 (3) 3-6(4) 2-6(6) 48 (6) 364 364 354 — _-
5 3-6 (4) 3-5 (4) 47(5) 49(6) 2-5(3) 2-5 (3) 2-5(3) 2406) — — —
6 3-7 (4) 2-4 (3) 2-5 (3) 2-6(3) 1,2 (@) 2 14 (2) 14@)
7 4-8 (5) 3-6 (4) 3-8 (5) 2-6(4) 2-7 (4) 2-6(3) 2-5 (3) 2,3 (3)
8 2-6 (3) — 2 1-5 (3) 1-5 (3) 1-40) 2-5 (3) 3-6 (4)
DF 2-5) (4) 12,()) | 2-6 (4) 1 l 1 1 ] —- —
LOW 2-916) == = 14 (1) 1403) 240) 2403) 2-5 (4) = — —
11 47 (4) = os = 1 1 2-5 (4) 2-5 (3)
12 — — = = = = = =
13 — — = = = = = = — —
base of gonocoxite. Claspette. Dorsal lobe
of claspette with 3 strong, closely appressed
setae, nearly equal in length, 2 setae insert-
ed nearly at the same level, rounded in lat-
eral view and 1 narrower seta inserted
slightly basal to the other 2 setae; ventral
lobe with 1 long, apical, slender seta; lateral
and mesal surfaces of claspette with nu-
merous small setae. Gonostylus with 10
minute setae on dorsal surface; gonostylar
claw short, spiniform and blunt. Aedeagus
with 5-8 pairs of leaflets, depending on po-
sition these appear as laminar, truncate at
apex or bluntly pointed; the most mesal leaf-
let about 0.45 of aedeagus length, uniform-
ly sclerotized, the other leaflets with the
outer edge less sclerotized; largest leaflets
with a few denticles toward apex and along
one or both edges, no basal denticles ap-
parent.
Pupa (Fig. 2).—Position and develop-
ment of setae as figured; range and modal
number of branches in Table 2. Integument
weakly pigmented with a mottled pattern of
dark pigmentation on wing case and leg
cases, antennal case darker on outer 0.5
with dark pigmentation at flagellomere
joints; integument near base of trumpet and
metathoracic wings with poorly defined
dark spots; paddle more darkly pigmented
on anterolateral half. Cephalothorax: Trum-
pet laticorn, tragus finger-like, slender, ta-
pered to apex. Secondary cleft absent. Ab-
domen: Terga and sterna II—VIII with nu-
merous small spicules, more evident mesal-
ly and posteriorly; lateral margins of terga
III-—VIII with numerous spicules, most
prominent on last segments. Setae 1-III—VII
and 5-IV—VII surrounded by strongly scler-
otized, spine-like protuberances, more
prominent on segments V—VII. Seta 9-II-—
VIII peglike to long and pointed without
aciculae. Segment VII 1.02—1.24 (mean
1.10) length of segment VI; segment VIII
1.13—1.55 (mean 1.41) length of segment
VI. Width/length (width at posterior mar-
gins) of segment VI 2.58—2.94 (mean 2.75),
VII 2.18—2.56 (mean 2.33), VIII 1.69—2.20
(mean 1.84). Paddle: Length 0.76—0.90 mm
(mean 0.81 mm), width 0.62—0.74 mm
(mean 0.68 mm), length/width 1.14—1.24
(mean 1.20); somewhat rounded, more
strongly pigmented on basolateral half; re-
fractile index 0.86—0.90 (mean 0.88); length
of marginal spicules 0.03—0.05 mm (mean
0.04 mm).
Larva (Fig. 3).—Position and develop-
ment of setae as figured; range and modal
VOLUME 99, NUMBER 3
3°20
‘iy
Fig. 3. Anopheles shannoni. Larva.
467
468
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 3. Larval setal branching for Anopheles shannoni: range, mode (). Most based on counts of twenty
setae. In the case of equal modes, the highest is given. A range shown without a mode indicates that the mode
was indefinite.
Thorax
Head ————————————————————
No. (e P M T
Abdominal Segments
I II I IV Vv
0 _ 1 = = 3-5) 4) See Oona
al 225 (Gyan e011) 2-574): » S29nG)Ga4 15-20 (19) 18-22 (20) 14-20 (20)
a “29 T-A2(8) 9226'(4) 225 (2) 42956) B= 12" (10). «6-1 (ye 228 (ee)
3 14-94°@8) a 1 FAA 2 g=4 (py 1 1,2 (1) DANA Sh)
A 4-74)" 59g deye=4 6) 4-7 (6) 7211 7) 6-8 ©). B54) (Fes a=)
5 197-26.(022):=30 I >30 42706) 714 0) 9710 4-6 (5) 48 (5)
6 16-24 (20) 1 2-5 (3) 3-5 (4) >30 >30 >25 2,3 (2) 232)
7 i7=24s(19) =30 3-7 |) >30 >30 >25 4-8(5) | B27 Gea
Be 236) >30 9-19 (10) >30 = 364) 24a 3,4 (4) 3-5 (5)
512 1 1 1 7213 (7), 10-15,15) 7-16 G0). 6-1 eo 1O ton
10: 123°@)" 1 I 1 L222) B=si4y. ADB) 131) ren AS)
i 20 1 1 1 3,4 (4) 2,3 (3) 2432). 325679246)
ie 2473). 1 1 DEAN(B)h 2-413) 2.390) 24.3). | /325vGi eee oka)
13 224 Gy 13226115) 52106) Asse) 7 16.(8) (G1ae ee 16 5-7)
iA 1-413) B-9@ ie 5219.12) aE: = = 4k” Ese ee = sas
15:73) = = _ = a =a = =
number of branches in Table 3. Head: An-
tennal length 0.29-0.33 mm (mean 0.31
mm), tapered toward apex, 5.05-—6.71
(mean 5.69) longer than wide; with spicules
longer and more numerous ventrally and in
vicinity of seta 1-A; dorsal surface with a
few spicules; seta 1-A with 7—13 branches,
inserted 0.26—0.32 (mean 0.30) from base
of antenna; seta 2-A pointed. Seta 2-C var-
ies from single and aciculate to strongly
aciculate or with up to 9 branches on apical
0.3, length 0.90—1.16 (mean 1.04) length
3-C, seta 2-C close to mate of opposite side,
distance between bases/width of base of
single seta 1.06—1.98 (mean 1.64); 3-C 18
to 24 branched (mode 18), clypeal index
(distance between bases of 2-C and 3-C on
one side/distance between the bases of 2-C)
1.67-2.79 (mean 2.21). Thorax: Seta 1-P
2-5 branched; setae 9-12-P single; 12-M
about 0.30 length of 9, 10-M; 11-M very
short, single; 3-T weakly developed, pal-
mate; 11, 12-T very short. Abdomen: Seta
1-I-VII palmate, seta 1-I,I1] weakly devel-
oped, leaflets broad, with jagged margins,
apex weakly pigmented; 9-I with 7-13
branches; 8—II with 3—6 branches; 6-IV,V
with 2,3 branches. Pecten with 15—20 teeth;
arrangement of teeth alternating long and
short, with 7-10 long and 7—11 short; long
spines 2.26—-3.93 (mean 3.17) length of
short spines. Seta 1-X not inserted on sad-
dle. Integument of posterior margin of seg-
ment X with numerous, strongly developed
spicules.
Material examined.—Holotype @ with
the following labels: handwritten ‘“‘shan-
noni’’; a printed red label “‘type no. 44166
USNM”; two printed white labels, one
‘Para, Braz. Apr. 1930, N.C. Davis’, an-
other ‘“‘animal bait’’. Paratypes, 2 2; same
as holotype (one marked by RW as proba-
bly a paratype did not have an original
paratype label). In addition, 49 2, 4 6, 13
larval exuviae, 15 pupal exuviae and 4 6
genitalia as follow. BRAZIL, Para State,
Belém City, Nova Timbo, 3 progeny broods
from females collected from human bait,
10.V. 1989 by J. B. Lima: BR 002(1), 1 2
2 Le 2 Pe; BR 002@), 2 2: 2:6 4\Leaike:
BR: 002@G); 7-2:1° Gig genes) Leo Re:
BR 002(X), 2 2; biting cow, 14.IV.194?,
Komp coll. and det., 1 2; Amazonas State,
Labrea, Rio Ituxi, Floresta, 18.1.1984, J.
Bento coll. and det., 1 d 1 ¢ genitalia; Ma-
naus, VI.1931, R.C. Shannon col. and det.,
VOLUME 99, NUMBER 3
Table 3. Extended.
Abdominal Segments
VI VII VIII xX
2-5 (4) 1-5 (4) 1-4 (2) =
15-22 (19), 13222-G7) °2,3:@2) 1
6-10 (7) 7-12 (8) 5-8 (6) 18-25 (20)
1-4 (2) 3-7 (5) 6-10 4-6 (6)
1 1-3 (2) I 9,10 (9)
6-9 (8) 7-11 (9) 4,5 (5) =
4-8 4-8 (6)
3-7 (3) 4-7 (4)
4,5 (5) 6-8 (7)
6-10 (10) 4-8 (5)
4-6 (5) 7-15 (11)
3-5 (3) 3
352) 2-4 (3) = am
6-11 (9) 4-7 (5) ae ee
2,3 (2) 1-3 (1) I -
3 2. PERU, Iquitos, III,IV.1931, R.C. Shan-
non coll., 15 2. GUYANA, Sector Malar.
Lab. TS&B.C., 19.VII.1944, T.K. Yolles
coll. and det. 1 2; on man, 1942, T.K. Yol-
les coll. and det., 16 2. SURINAM, Para-
maribo, biting man, 3.IX.1943, D.G. Hall
Coley oe
Annotated bibliography.—Shannon 1933:
136 (6; 2* [genitalia, marginal wing
scales, halter]; pupa* [trumpet, cephalotho-
rax, abdomen]. Iquitos, Peru; Belém and
Manaus, Brazil); Vargas 1942: 72 (2, key);
Russell et al. 1943: 49 (2, key. British Gui-
ana); Cerqueira 1943: 18 (Beni and Terr. de
Colonias, Bolivia); Causey et al. 1944: 3
(egg*, key); Causey et al. 1946: 26 (d*
[claspette, aedeagus], key); Deane et al.
1946a: 13, (2 * [wing, cerci, hindtarsomere
1], key); Deane et al. 1946b: 37, 41 (larva*
[setae 2,3,4-C, antenna, setae 1,2,3-P], key.
Mato Grosso, Amazonas and Para, Brazil);
Deane et al. 1948: 917 (distribution map,
northern Brazil); Correa 1950: 81 (2 and
larva in keys); van der Kuyp 1950: 63 (2,
36 and larva, in keys. Moenga, Surinam);
Lane 1953: 204 (¢* [wing], key; 6; pupa*
[trumpet, abdomen]; larva* [illus. from
Deane et al. 1946b]; egg* [illus. from Cau-
469
sey et al. 1944]); Vargas 1959: 385 (¢ gen.
in key); Forattini 1961: 172, 181, 186 (¢,
3 gen. and larva in keys); Forattini 1962:
342 (3 gen.* [claspette, aedeagus], key; @
and larva in keys); Garcia and Ronderos
1962: 149 (¢* [wing], key; ¢ gen. in key;
larva* [setae 2,3-C, 6-IV,V], key); Gorham
et al. 1967: 25, 40, 61; (2 and larva in
keys); Morales-Ayala 1971: 138 (Loreto,
Peru); Rambajan 1987: 149 (Guyana).
Distribution.—Reported from northern
Amazonian Brazil, Guyana, eastern Peru,
Bolivia and Surinam. From this distribution
it can be assumed to also occur in Ama-
zonian Ecuador, Colombia and Venezuela.
Biology.—Little is known about the bi-
ology of adult and immature stages of An.
shannoni. The type specimens were cap-
tured with animal bait inside the jungle dur-
ing morning and evening hours. Adults
were observed to be zoophilic since they
were rarely found in houses, but commonly
collected in animal shelters. Adults were
also collected with a Shannon trap in the
Amazon forest. Larvae were reported from
forest ponds or pools (Deane et al. 1946b,
Shannon 1933), from stagnant river waters
and small shaded streams, and in flooded
forests with clear water generally full of de-
composing leaves, among shrubs and tree
trunks (Deane et al. 1948). Deane et al.
(1948) also reported that the larvae were
found associated with An. (Nyssorhynchus)
darlingi Root and An. (Ano.) mediopunc-
tatus (Theobald) and that adults seldom en-
tered houses and fed mostly on horses at
dusk.
Discussion.—In comparison to other spe-
cies in the Arribalzagia Series (subgenus
Anopheles) (Reid and Knight 1961, Wilk-
erson and Peyton 1990), the adult female
An. shannoni is similar to An. minor Da
Costa Lima, An. peryassui Dyar and Knab
and An. mattogrossensis Lutz and Neiva in
having mostly dark-scaled tarsi with narrow
rings of pale scales at the tarsomere artic-
ulations. However, the former two species
have posterolateral abdominal scale tufts
and speckled tibiae and femora, while An.
470
peryassui and An. mattogrossensis have
neither. In many other respects An. shan-
noni is quite similar to An. minor but An.
shannoni has wide wing scales, the apical
half of the cercus is strongly constricted and
there is no speckling on hindtarsomere 1.
Anopheles minor has narrow wing scales, a
rounded cercus and speckling on hindtar-
somere 1.
The male genitalia of An. shannoni has
the ninth tergal lobes short and triangular
while they are short to long but rounded in
the other three species. Anopheles shannoni
has 5-8 large subequal aedeagal leaflets
with one edge usually thinner and nearly
transparent, and with small apical denticles.
Anopheles minor and An. mattogrossensis
both have one pair of leaflets (An. minor
also has several very small pairs) and An.
peryassui has 4-5 pairs of uniformly scler-
otized leaflets without denticles. Of the four
species only An. shannoni has a single pri-
mary seta on the ventral lobe of the clas-
pette.
In the larval stage An. shannoni can be
distinguished from other An. (Anopheles)
by the following combination of characters:
seta 2-A lanceolate with a fine apical fringe
on one margin; seta 3-C with 18—24
branches; seta 1-P short with 2—5 branches;
setae 9-12-P single; seta 6-IV,V 2-3
branched and; the pecten plate with alter-
nating short and long teeth.
In the pupal stage, An. shannoni can be
easily recognized by setae 1-III—VII and
5-IV—VII which have bases surrounded by
spinelike projections. Also, the trumpet has
a slender, fingerlike tragus, the secondary
cleft is absent and the paddle is strongly
pigmented on the basolateral half.
The egg is unusual in having numerous
frills (Causey et al. 1944, Lounibos et al. in
press), similar to An. peryassui Dyar and
Knab (Causey et al. 1944, Linley and Loun-
ibos 1994).
ACKNOWLEDGMENTS
We thank T. R. Litwak for Figs. 1 and 3
and E. Roberts for the pupal drawings on
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. J. B. Lima was responsible for the
collection and rearing of specimens from
Ilha de Maraj6, Para, Brazil, that provided
the stimulus for this work. E. L. Peyton fur-
nished a critical review of an early version
of the manuscript. Partially supported by
Grant no. 95/7157-2 from the Fundacao de
Amparo a Pesquisa do Estado de Sao Paulo
(FAPESP), Brazil
LITERATURE CITED
Belkin, J. N., S. J. Heinemann, and W. A. Page. 1970.
Mosquito studies (Diptera, Culicidae). XXI. The
Culicidae of Jamaica. Contributions of the Amer-
ican Entomological Institute 6(1): 1-458.
Causey, O. R., L. M. Deane, and M. P. Deane. 1944.
An illustrated key to the eggs of thirty species of
Brazilian anophelines, with several new descrip-
tions. American Journal of Hygiene 39(1): 1-7.
Causey, O. R., L. M. Deane, and M. P. Deane. 1946.
Studies on Brazilian anophelines from the north-
east and Amazon regions. II. An illustrated key by
male genitalic characteristics for the identification
of thirty-four species of Anophelini from the
northeast and Amazon regions of Brazil, with a
note on dissection technique. American Journal of
Hygiene Monographic Series 18: 21-31.
Cerqueira, N. L. 1943. Lista dos mosquitos da Bolivia
(Diptera, Culicidae). Memorias do Instituto Os-
waldo Cruz 39: 15-36.
Correa, R. R. 1950. Alguns informes sébre Anapheles
(sic.) (Arribalzagia) intermedius (Chagas, 1908),
(Diptera, Culicidae). Arquivos de Higiene e Saude
Publica 24(39—42): 79-89.
Davis, N. C. 1931. A new anopheline mosquito from
Para, Brazil. American Journal of Hygiene 8:
345-348.
Deane, L. M., O. R. Causey, and M. P. Deane. 1946a.
Studies on Brazilian anophelines from the north-
east and Amazon regions. I. An illustrated key by
adult female characteristics for the identification
of thirty-five species of Anophelini, with notes on
the malaria vectors (Diptera, Culicidae). American
Journal of Hygiene Monographic Series 18: 1—20.
Deane, M. P., O. R. Causey, and L. M. Deane. 1946b.
Studies on Brazilian anophelines from the north-
east and Amazon regions. III. An illustrated key
by larval characteristics for the identification of
thirty-two species of Anophelini with descriptions
of two larvae. American Journal of Hygiene Mon-
ographic Series 18: 35—50.
Deane, L. M., O. R. Causey, and M. P. Deane. 1948.
Notas s6bre a distribuicao e a biologia dos ano-
felinos das regides nordestina e amaz6nica do
Brasil. Revista do Servigo Especial de Saude Pub-
lica 1(4): 828-965.
VOLUME 99, NUMBER 3
Forattini, O. P. 1961. Chaves para a identificacao do
genero Anopheles Meigen, 1818, da regiao neo-
tropical (Diptera, Culicidae). Revista Brasileira de
Entomologia 10: 169-187.
Forattini, O. P. 1962. Entomologia Medica. Vol 1. Par-
te Geral, Diptera, Anophelini. Faculdade de Hig-
iene e Saude Publica, 662 pp.
Garcia, M. and R. A. Ronderos. 1962. Mosquitos de
la Republica Argentina. I. Tribu Anophelini (Dip-
tera-Culicidae-Culicinae). Buenos Aires (Prov.)
Comision para Investigaticiones. Cientificas. (?) 3:
103-212.
Gorham, J. R., C. J. Stojanovich, and H. G. Scott.
1967. Clave ilustrada para los mosquitos anofel-
inos de Sudamerica oriental. U.S. Department of
Health Education and Welfare, 64 pp.
Harbach, R. E. and K. L. Knight. 1980. Taxonomists’
Glossary of Mosquito Anatomy. Plexus Publish-
ing, Inc., Marlton, New Jersey, 413 pp.
Harbach, R. E. and K. L. Knight. 1982. Corrections
and additions to Taxonomists’ Glossary of Mos-
quito Anatomy. Mosquito Systematics (1981) 13:
201-217.
Kuyp, E. van der. 1950. Contribution to the study of
the malarial epidemiology in Surinam. Konink-
lijke Vereeniging Indisch Instituut 89: 1-146.
Lane, J. 1953. Neotropical Culicidae. 2 volumes. The
University of Sao Paulo, 1112 pp.
Linley, J. R. and L. P. Lounibos. 1994. The remark-
able egg of Anopheles peryassui (Diptera: Culic-
idae). Mosquito Systematics 26: 25-34.
Lounibos, L. P., D. Duzak, J. R. Linley, and R. Lou-
renco de Oliveira. In press. Egg structure of
Anopheles fluminensis and Anopheles shannoni.
Memorias do Instituto Oswaldo Cruz.
Morales-Ayala, F 1971. A list of the mosquitoes of
Peru. Mosquito Systematics Newsletter 3: 138—
145.
471
Rambajan, I. 1987. An annotated checklist of Guyana,
South America. Mosquito Systematics 19: 146—
161.
Reid, J. A. and K. L. Knight. 1961. Classification
within the subgenus Anopheles (Diptera, Culici-
dae). Annals of Tropical Medicine and Parasitol-
ogy 55: 474-488.
Reinert, J. E 1975. Mosquito generic and subgeneric
abbreviations (Diptera: Culicidae). Mosquito Sys-
tematics 7: 105-110.
Russell, P. E, L. E. Rozeboom, and A. Stone. 1943.
Keys to the anopheline mosquitoes of the world.
With notes on their distribution, biology and re-
lation to malaria. American Entomological Soci-
ety. Lancaster Press. 152 pp.
Shannon, R. C. 1933. Anophelines of the Amazon
Valley. Proceedings of the Entomological Society
of Washington 35: 117-143.
Vargas, L. 1942. Las hembras americanas del subge-
nero Anopheles (Dit, Culicidae, Anopheles). Re-
vista del Instituto de Salubridad y Enfermedades
Tropicales 3: 67-74.
Vargas, L. 1959. Lista de Anopheles de las Americas
y su identificacion por caracteres masculinos. Re-
vista del Instituto de Salubridad y Enfermedades
Tropicales 19: 367-386.
Wilkerson, R. C. 1990. Redescriptions of Anopheles
punctimacula and An. malefactor (Diptera: Culic-
idae). Journal of Medical Entomology 27: 225-—
247.
Wilkerson, R. C. 1991. Anopheles (Anopheles) cald-
eroni n.sp., a malaria vector of the Arribalzagia
Series from Peru (Diptera: Culicidae). Mosquito
Systematics 23: 25-38.
Wilkerson, R. C. and E. L. Peyton. 1990. Standard-
ized nomenclature for the costal wing spots of the
genus Anopheles and other spotted-wing mosqui-
toes (Diptera: Culicidae). Journal of Medical En-
tomology 27: 207-224.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 472-476
A NEW GENUS AND SPECIES OF ALLANTINAE (HYMENOPTERA:
TENTHREDINIDAE) FROM SOUTHEASTERN UNITED STATES
DAVID R. SMITH AND TERENCE L. SCHIEFER
(DRS) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S.
Department of Agriculture, c/o National Museum of Natural History, MRC 168, Wash-
ington, DC 20560, U.S.A.; (TLS) Mississippi Entomological Museum, Box 9775, Mis-
Sissippi State, MS 39762, U.S.A.
Abstract.—Allantunicus autumnalis, a new genus and a new species of Tenthredinidae,
subfamily Allantinae, is described from the sand dune area of coastal Alabama. The genus
and species are diagnosed and compared to other North American Allantinae. Evidence
strongly suggests that the larva feeds on inflorescences of Polygonella gracilis (Polygon-
aceae).
Key Words:
The specimens on which the following
new genus and new species are based were
collected in autumn from Polygonella grac-
ilis (Nutt.) Meisn. (Polygonaceae) in the
sand dune area of coastal Alabama. Obser-
vations from separate larval collections in
this same area strongly suggest that the lar-
va feeds on the inflorescences of Polygo-
nella gracilis, a plant of the sandhills of the
Coastal Plain of the southeastern United
States. A single specimen was initially dis-
covered in the collection of the Mississippi
Entomological Museum. In October 1996,
the junior author visited the same collecting
area and obtained additional specimens, in-
cluding males. The new genus and species
is morphologically distinct from all other
North American Allantinae, and its pre-
sumed habits are unique among the Allan-
tinae. Some of the remaining sawflies to be
discovered in the Nearctic Region no doubt
live in restricted habitats and are active dur-
ing an unusual time of the season. The pos-
sibility of this being an adventive species
was explored; however, there are no Neo-
tropical allantines similar to this species and
sawfly, Tenthredinidae, Allantinae, Polygonella gracilis, Polygonaceae
no genus and species comparable to it from
the rest of the world.
Allantunicus Smith, new genus
Type species.—Allantunicus autumnalis
Smith, new species.
Antenna (Fig. 4) filiform; Ist and 2nd
segments each longer than broad; 3rd seg-
ment about 1.7 X longer than 4th segment.
Head narrowing behind eyes in dorsal view,
distance behind eyes equal to half visible
eye length; postocellar area as broad as
long; mandibles symmetrical, each biden-
tate; malar space nearly linear; clypeus
shallowly emarginate, laterally acute with-
out rounded lobes; short genal carina indi-
cated only below eye near mandible. Pro-
pleura broadly rounded and meeting on
meson; mesopleuron and mesonotum
smooth, without punctures. Tarsal claw
(Fig. 3) with small inner tooth at center of
claw, shorter than outer tooth; basal lobe
absent. Abdomen without pairs of white
spots on dorsum. Forewing with anal cross-
vein oblique, with 4 cubital cells, M and Rs
+ M meeting Sc + R at same point. Hind-
VOLUME 99, NUMBER 3
wing without cells Rs and M;; length of pet-
iole of anal cell about half width of cell.
Etymology.—The name is based in part
on the subfamily name, Allantinae, and the
Latin unicus, indicating the unusual and
unique characteristics of the genus.
Discussion.—The genus is separated
from other allantine genera by the follow-
ing combination of characters: short genal
carina present only near mandible below
eye; shallow clypeal emargination with
acute lateral lobes; smooth, shining thorax
without punctures on mesepisternum; short
inner tooth of the tarsal claw; lack of closed
cells Rs and M in the hindwing; and third
antennal segment longer than fourth seg-
ment. Allantunicus will key to Somanica
Smith in couplet 12 of my key to genera
(Smith 1979). Somanica differs from Allan-
tunicus by the subequal third and fourth an-
tennal segments, lack of a genal carina,
rounded lateral lobes of the clypeus, and
long inner tooth of the tarsal claw, nearly
as long as the outer tooth. In addition char-
acteristics of the sheath and lancet (Figs. 1,
2, 5) of Allantunicus are unique among Ne-
arctic Allantinae, namely the emarginate
apex of the sheath in lateral view, slight
widening of the sheath toward the apex in
dorsal view, and presence of ctenidia on the
annuli of the lancet.
Allantunicus belongs to the tribe Empri-
ini, as defined by Smith (1979): mandibles
symmetrical, bidentate; clypeus shallowly
emarginate, propleura meeting broadly on
the meson; mesopleuron without punctures;
and veins M and Rs+M meeting Sc+R at
the same point in the forewing.
This does not appear to be an adventive
taxon. A survey of material in the National
Museum of Natural History, Smithsonian
Institution, and use of Malaise’s (1963) key
to world genera indicated no similarities to
exotic taxa. Allantunicus runs to couplet
106 in Malaise’s key which identifies Mal-
lachiella Malaise (from Burma and _ the
Himalayas) and Heptapotamius Malaise
(from Turkestan). Mallachiella differs by
the presence of a basal lobe on the tarsal
473
claw, a compressed antennal flagellum, sub-
symmetric mandibles, and a very deeply,
semicircularly incised clypeus with long,
acute lateral lobes. Heptapotamius differs
by the head strongly enlarged behind the
eyes in dorsal view, and the clypeus incised
for about a quarter of its media! length and
with blunt lateral teeth.
Allantunicus autumnalis Smith,
new species
(Figs. 1-7)
Female.—Length, 8.5—9.0 mm. Antenna
and head black; mandible dark orange with
apex blackish, clypeus whitish; basal 3
maxillary palpal segments and labial sub-
mentum orange, rest of palpi and labium
blackish. Thorax orange with anterior mar-
gin of propleuron, entire prosternum and
mesosternum, and spot on lower portion of
metepimeron, black. Abdomen orange;
sheath black. Legs orange with apical 3
midtarsal segments and entire hindtarsus
blackish. Wings moderately, uniformly in-
fuscated; veins and stigma black.
Antenna as in Fig. 4, length 1.6 head
width. Eyes slightly converging below;
lower interocular distance subequal to eye
length. Hindbasitarsus shorter than remain-
ing hindtarsal segments combined, ratio
1.8:3.0. Head and body entirely shining and
impunctate. Sheath (Figs. 1, 2) in lateral
view with posterior margin slightly con-
cave, in dorsal view slightly expanded to-
ward apex with indication of small scopa;
with long hairs curved mesally. Lancet as
in Fig. 5, with stout ctenidia on annuli; ser-
rulae low, only apical 7 with fine posterior
subbasal teeth.
Male.—Length, 8.0—9.0 mm. Coloration
similar to female. Genitalia as in Figs. 6, 7.
Holotype-—Female from Alabama, la-
beled ‘“‘Ala., Baldwin Co., Gulf Shores,
30°16'47” N, 87°42'49" W, 16 Oct. 1996,
T.L. Schiefer,”’ ‘‘on flowers of Polygonella
gracilis in dredge spoil dunes,’’ Deposited
in the National Museum of Natural History,
Smithsonian Institution, Washington D.C.
on indefinite loan from the Mississippi En-
474 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-7. Allantunicus autumnalis. 1, Sheath, lateral view. 2, Sheath, dorsal view. 3, Tarsal claw. 4, Antenna.
5, Lancet. 7, Male genitalia, genital capsule, left side, ventral view. 7, Male genitalia, penis valve, lateral view.
VOLUME 99, NUMBER 3
tomological Museum, Mississippi State
University.
Paratypes—ALABAMA: Same data as
holotype (4 2); same data but 17 Oct. 1996
(3 2); same data but 14 Oct. 1991 (1 @);
same data but 17 Oct. 1996, in flight near
Polygonella gracilis in dredge spoil dunes
Giecd): ssamewdata. buts 17, Oct») 1996;
30°16'52” N, 87°42'03” W, on flowers of
Polygonella gracilis in dredge spoil dunes
(2 2, 1 3). Deposited in the Mississippi
Entomological Museum, Mississippi State
University; National Museum of Natural
History, Smithsonian Institution, Washing-
ton, D.C.; and Canadian National Collec-
tion, Ottawa.
Etymology.—The name is derived from
the autumn flight of the species.
Host.—Polygonella_ gracilis (Nutt.)
Meisn., wireweed (Polygonaceae) (see dis-
cussion).
Color variation.—Several specimens dif-
fer in color from the typical form as de-
scribed. The mesosternum and mesepister-
num of two females are mostly black except
for the orange upper third of the mesepi-
sternum and most of the metepisternum,
cervical sclerites, metapleuron, and base of
midcoxa. One male has a black mesoster-
num and mesepisternum except for a small
orange spot on the upper corner of the me-
sepisternum, the metepleuron, and most of
the lateral lobes of the mesonotum.
Discussion.—On 16-17 October 1996,
the junior author collected most of the type
series from Polygonella gracilis in a sand
dune area at Gulf Shores, Alabama. All
were net-collected on flowers or in flight
near the plants. The 11 specimens collected
on flowers were always oriented in align-
ment with the axis of the flower cluster.
They were found on both horizontal and
vertical flower clusters with their head fac-
ing either toward or away from the main
stem of the plant. All except two of these
individuals were sitting motionless on the
flowers. One female observed in late after-
noon occasionally turned around on the
flower cluster to face the opposite direction.
475
It appeared to be tapping the flowers with
its antennae, but no oviposition was ob-
served. The next morning another individ-
ual was observed flying from one flower
cluster to another on the same plant and
then turning around several times like the
one observed the previous afternoon. Two
individuals were on flower clusters in
which most of the flowers had not yet
opened. The five specimens collected in
flight that morning flew among the Poly-
gonella in a fairly slow and erratic zigzag
pattern. However, their evasive flight was
very fast and direct. After about 10:30 am,
there appeared to be a marked decrease in
sawfly activity, and no additional specimens
were observed at the type locality. This cor-
responded to an increase in temperature and
increased pollinator activity. However,
three additional specimens were collected
about noon in similar habitat 0.6 miles from
the original site. Only males were caught in
flight; all females and one male were taken
on the flower clusters.
Polygonella gracilis grows in the pure
white sand characteristic of the dune habi-
tats in the vicinity of Gulf Shores. The fore-
dune and backdune habitats close to the
coast would seem to be suitable habitats for
the sawfly since P. gracilis is present. But
despite several Malaise traps set for two
days in these habitats and much visual
searching, no sawflies were found there.
The sandy areas where the sawflies were
collected are along the north side of the In-
tracoastal Waterway where P. gracilis is
more abundant. This habitat is apparently
man-made having been created through the
dumping of spoil from the dredging of the
waterway. The area has subsequently been
colonized by vegetation more characteristic
of the foredune and backdune habitats that
lie several miles closer to the coast. Ac-
cording to Paul Warren, Army Corps of En-
gineers, Mobile District, Irvington Office
(personal communication), the first channel
for this section of the waterway was dug in
1934 and then enlarged in 1944. This prob-
ably accounts for much of the dredge spoil
476
that created the sawfly habitat. However,
since that time the waterway has continued
to be dredged periodically, most recently in
the mid-1980’s. The habitat at the type lo-
cality has recently been greatly altered by
the removal of much of the sand. Most of
the remaining habitat suitable for the sawfly
lies in a narrow band at the south end of
the spoil area.
This is probably the species discovered
in the larval stage by W. McDearman, Mu-
seum of Natural Sciences, Jackson, Missis-
sippi. According to McDearman (1990 in
litt.) when he was at Gulf Shores, Alabama,
in early November botanizing along the
dunes, ‘‘I collected what appears to be saw-
fly larvae which were exclusively feeding
on Polygonella gracilis (Nutt.) Meisn. (wire-
weed). The flowering period for these
plants is a bit unusual—late summer and
fall—from September through November.
As with Rhadinoceraea [see Smith and
McDearman 1990]—flowers were the pri-
mary food. These plants are indeed very
slender and wiry with very little stem cor-
tex (stem diameters rarely over 5 mm). I
did not observe any indications that larvae
emerged from eggs laid within the stem. I
watched one larva disperse over a period of
about an hour—but didn’t see it burrowing
in the soil to pupate. No adults were seen.”’
In a subsequent letter, McDearman (1991 in
litt.) stated that he returned to the coast in
September 1990 hoping to find some larvae
and adults on Polygonella gracilis, but was
too early as few plants were in flower.
Adults were collected near the area
McDearman made these observations, from
Polygonella gracilis flowers, and were col-
lected late in the year (mid-October) during
the flowering time of the host (larvae were
observed in November by McDearman).
These pieces of circumstantial evidence
strongly suggest that larvae of A. autum-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
nalis feed on the inflorescences of Polygo-
nella gracilis.
Polygonella gracilis is an annual of sand-
hills and occurs in the Coastal Plain from
South Carolina to Florida west to Louisiana
(Rickett 1967). The sawfly may have a
much wider distribution but has escaped
discovery because of its restricted habitat
and late seasonal occurrence which is very
unusual for sawflies.
ACKNOWLEDGMENTS
W. McDearman provided information on
larval observations. Figure 5 was drawn by
Linda Lawrence, staff artist, Systematic En-
tomology Laboratory, USDA. Cathy An-
derson, Museum Technician, Systematic
Entomology Laboratory, USDA, assisted
with the final draft of the manuscript. The
contribution by T.L. Schiefer was approved
for publication as Journal Article No.
J-9121 of the Mississippi Agricultural Ex-
periment Station, Mississippi State Univer-
sity, with support from State Project MIS-
6538. I thank the following for reviewing
the manuscript: Henri Goulet, Agriculture
Canada, Ottawa; Nathan N. Schiff, USDA,
Stoneville, Mississippi; and N.E. Woodley
and A.S. Menke, Systematic Entomology
Laboratory, USDA, Washington, DC.
LITERATURE CITED
Malaise, R. 1963. Hymenoptera Tenthredinoidea, sub-
family Selandriinae, key to the genera of the
world. Entomologisk Tidskrift 84: 159-215.
Rickett, H. W. 1967. Wild Flowers of the United
States. Volume Two, Part One. The Southeastern
States. McGraw-Hill Book Company, New York.
322 pp.
Smith, D. R. 1979. Nearctic sawflies IV. Allantinae:
adults and larvae (Hymenoptera: Tenthredinidae).
United States Department of Agriculture, Techni-
cal Bulletin Number 1595, 172 pp. + 24 plates.
Smith, D. R. and W. McDearman. 1990. A new Rhad-
inoceraea (Hymenoptera: Tenthredinidae) feeding
on Zigadenus (Liliaceae) from southeastern Unit-
ed States. Entomological News 101: 13-19.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 477-482
TWO NEW SPECIES OF ASYNDETUS LOEW, AND NOTES ON THE
IDENTITY OF A. INTERRUPTUS LOEW (DIPTERA: DOLICHOPODIDAE)
HAROLD ROBINSON AND MARK DEYRUP
(HR) Department of Botany, National Museum of Natural History, Smithsonian Insti-
tution, MRC 166, Washington, DC 20560, U.S.A.; (MD) Archbold Biological Station,
PO. Box 2057, Lake Placid, FL 33862, U.S.A.
Abstract.—Asyndetus archboldi is described as new from the sand hills of peninsular
Florida; A. wirthi is described from Jamaican and Antiguan material previously deter-
mined as A. interruptus. Observations are given on the type specimen of A. interruptus
Loew in Havana, Cuba.
Key Words:
Two new species of Asyndetus Loew
are described, the first based on collec-
tions from the xeric uplands of Florida,
and the second based on West Indian ma-
terial previously identified as A. interrup-
tus Loew.
The Florida Asyndetus is described and
discussed as part of an effort to catalog the
species of arthropods found in the Florida
Scrub habitat. Florida scrub is a fire-main-
tained habitat that occurs on xeric sand
ridges in scattered localities in Florida. The
scrub habitat supports a number of species
of plants and animals found nowhere else,
and the rapid disappearance and degrada-
tion of the remaining areas lends an urgen-
cy to the project of cataloguing the resident
arthropods. For more detailed descriptions
of Florida scrub, see Myers (1990) and
Abrahamson et al. (1984); for more infor-
mation on Florida scrub arthropods, see
Deyrup (1990) and Deyrup and Eisner
(1993).
All specimens are presently deposited
at either the National Museum of Natural
History, Smithsonian Institution, (USNM)
or the Archbold Biological Station (ABS).
Asyndetus, new species, Dolichopodidae, Florida, West Indies
Asyndetus archboldi Robinson and
Deyrup, new species
(Figs. 1-3)
Holotype male.—Length 1.9 mm; wing
1.6 mm by 0.6 mm. Face 1.4 times longer
than wide, sides parallel; face and front
with dense silvery pollen. Palpus black,
with several stout black setae; proboscis
black; upper postocular setae black, lower
postocular setae white. Antenna black,
scape widened apically, without hairs or se-
tae; pedicel about twice as long dorsally as
ventrally in external lateral view, with a
dorsal apical enlarged black seta; Ist fla-
gellomere with a long, dorsal, sub-basal
arista, encircled by a deep groove just dis-
tad of arista, so flagellomere appears divid-
ed into 2 parts.
Thorax dark gray with light gray pollen,
with extremely faint greenish and coppery
reflections when viewed in fluorescent
light, with both green and coppery reflec-
tions conspicuous in incandescent light; all
setae black; only 4 acrostichals in single ir-
regular row, 4 pairs of dorsocentrals; 2
large scutellar bristles; 1 black proepisternal
seta above fore coxa.
478
Coxa black, front and middle coxa with
an anterolateral irregular row of bristles,
hind coxa with an external basal bristle;
femora black with conspicuous green re-
flections and yellowish apices; front and
middle tibiae yellowish, hind tibia and all
tarsi black; hairs and setae on legs black;
middle and front tibiae with small seriate
setae and single enlarged apical seta, hind
tibia with paired anterodorsals and poster-
odorsals at basal fifth, at two-thirds and
subapically; an unpaired anterodorsal at ba-
sal third. Front basitarsus with a subapical
external cluster of small thick setae; 2nd,
3rd and 4th segments compressed, 5th seg-
ment depressed; pulvilli about as long as
rest of 5th segment, claws about half as
long as pulvilli, shorter than 3 apical bris-
tles of 5th segment; lengths of tarsal seg-
ments from base: tarsus I, 0.30, 0.09, 0.06,
0.04, 0.05 mm, pulvilli 0.05 mm; tarsus II,
0.35, 0.16, 0.11, 0.08, 0.08 mm, pulvilli
0.03 mm; tarsus III, 0.25, 0.18, 0.12, 0.08,
0.08 mm, pulvilli 0.02 mm.
Wings appearing rather smoky due to
dense, uniform microtrichae; veins dark
brown; costa ending at apex of 3rd vein;
4th vein discontinuous, the “‘broken’’ ends
apparently overlapping, at least as folds; ca-
lypter, its setae, and knob of halter, ivory.
Abdomen dark gray with thin gray pol-
len, with greenish and coppery reflections
that are almost imperceptible in fluorescent
light, conspicuous in incandescent light.
Tergal setae black, suberect, apical band on
each tergite larger; genital capsule with 4
large equal setae.
Allotype female.—Length 2.2 mm; wing
2.0 mm by 0.7 mm. Coloration, pollinosity,
wing venation, tibial setae similar to male.
Front tarsi not modified, setae of tergites
shorter and more decumbent.
Holotype.—Male: FLORIDA: Highlands
Co., Archbold Biological Station, 26 April
1996, M. Deyrup. On open sand at edge of
fire lane, Florida scrub habitat (USNM).
Paratypes—(USNM): 6 6, 3 2 (includ-
ing allotype): same locality, date, collector,
habitat as holotype; 1 2: same locality, col-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
lector as holotype, 30 Apr. 1989, road 6,
scrubby flatwoods; 2 2: same locality, col-
lector as holotype, 23 June 1989, white
sand trail, scrub; 1 2: same locality, collec-
tor as holotype, 29 June 1989, white sand
fire lane: 16, 1 9: 4 65/15 2svsamecto-
cality, collector as holotype, 16 June 1992,
inopina scrub, line road; 1 6, 8 2: same
locality, collector as holotype, 18 June
1992; 3 5, 3 2: same locality, collector as
holotype, 19 June 1992; 1 ¢: same locality,
collector, habitat as holotype, 18 Apr. 1996;
3 3, 7 2: same locality, collector, habitat
as holotype, 23 Apr. 1996; 2 6, 1 2: same
locality, collector, habitat as holotype, 24
Apr. 1996; 1 ¢: Lake Placid, Placid Lakes
Development, 15 June 1992, M. Deyrup;
same locality as holotype, 26 June 1989, R.
Shumate, Malaise trap, Ceratiola area.
(ABS): 9 3, 7 2: same locality, collector,
habitat as holotype, | May 1996.
Etymology.—Named for the Archbold
Biological Station, where most of the type
material was collected.
Diagnosis.—Because of the pale fore tib-
ia, black hind tibia, and the male face that
is higher than wide, Asyndetus archboldi
keys in Robinson (1964) to couplet 6, but
differs from all the species in that part of
the key by the small first flagellomere of
the antenna. Relationship might be closest
to some western species, particularly A. /a-
tus Van Duzee, from Bill Williams Fork,
Arizona, having a similar densely silvery
pollinose face and black palpi. The latter
species is distinctly larger, ca. 3.5 mm long,
is evidently from near water, and resembles
other possibly related western species in
having all the tibiae partly or wholly yellow
(Van Duzee 1916, 1919).
Discussion.—Florida scrub habitat is the
antithesis of habitats preferred by most Dol-
ichopodidae: it is comparatively sterile, xer-
ic, without nearby permanent water, with
clumps of sclerophyllous, drought-adapted
plants, and patches of bare, white sand.
Other species of Asyndetus are usually
found on the seacoast or around permanent
fresh water (Cole 1969). However, A. arch-
VOLUME 99, NUMBER 3
boldi shares a habitat trait with some of its
congeners: it is associated with sandy plac-
es. This is somewhat parallel to the situa-
tion in pygmy mole crickets (Tridactylidae)
which live on algae growing in wet, sandy
areas, except for a newly described species
from Florida scrub (Deyrup and Eisner
1996). This latter pygmy mole cricket feeds
on a recently discovered layer of algae that
grows about 3 mm below the surface of the
bare sand, where light is easily transmitted
through the translucent grains and where at-
mospheric drying is much reduced. The lar-
vae are known for only one species of
Asyndetus (Williams 1938), that naturally
occurs at the entrances of crab holes on
sandy beaches, and whose larvae have been
raised in vials where they were fed on dro-
sophilid maggots. Almost all dolichopodid
larvae are similarly predacious, but the an-
imals on which they feed occur in a wide
variety of habitats. At least some dolicho-
podid larvae are associated with algal mats
(Oldroyd 1964). Thus, we speculate that the
larvae of A. archboldi may be feeding in
the algal layer under the sand on which the
adults are found. The limited literature on
dolichopid larvae and pupae (Willams
1938; Smith 1952; Dyte 1959; Robinson
and Vockeroth 1981) would be augmented
by discovery and description of the imma-
ture stages of A. archboldi.
Adults of Asyndetus archboldi occur on
open sand at edges of fire lanes and in bare
patches between clumps of scrub plants.
These flies move quickly from place to
place on the sand, seldom flying more than
a few mm above the surface. They may
perch for a few minutes on debris, especial-
ly the small dead leaves of scrub oaks.
When individuals meet, they usually per-
form a rapid gyrating flight around each
other, then separate. A fly can be captured
by quickly placing a plastic bag over it,
then chasing it up into the bag, where it can
be sucked up with an aspirator. We have
collected one specimen in a Malaise trap,
but there seems to be no method for easily
capturing large numbers of specimens at
479
one time. We expect that, when dipterists
begin to scan the surface of open sand for
these tiny flies, that this species or similar
species will be found in scrub elsewhere in
Florida, and perhaps outside of the state as
well.
Asyndetus wirthi Robinson,
new species
(Figs. 4—6)
Holotype male.—Length 4.0 mm; wing
3.5 mm by 1.2 mm. Face ca. %4 to ¥; as wide
as high, not recessed, white pollinose; front
not noticeably wider than face at antennae,
metallic greenish color almost totally ob-
scured by grayish-white pollen. Palpus and
proboscis black, the former with numerous
black setae. Antenna black; pedicel only
slightly produced above and on inner side,
not longer above than below; first flagello-
mere scarcely as long as high, with very
short, blunt point below arista, arista borne
at distal %4 of upper edge, very near tip.
Lower postocular setae white.
Thorax mostly metallic green obscured
with yellowish-gray pollen, with rather dis-
tinct brown pollinose stripe dorsally be-
tween rows of dorsocentrals; acrostichals
distinctly biseriate; scutellum with small
hair on lateral margin; with series of 5 or 6
small proepisternal setae above fore coxa of
which those nearest the coxa are longer.
Legs dark, with some yellow at knees;
coxae and femora metallic green with gray-
ish pollen. Setae on anterior surfaces of
coxae coarse, black. All femora with some-
what irregular series of anteroventral and
posteroventral setae, half to three-fourths as
long as widths of femora, a stouter antero-
ventral seta at tip of hind femur, 2 or 3 lon-
ger setae anterodorsally near tip of hind fe-
mur. Fore tibia with 2 or 3 anterodorsals,
third and largest anterodorsal near middle,
1 distinct posterodorsal near middle, 2
small apicals above and below; middle tibia
with 2 large anterior bristles near /; and %,
small anterodorsal near 7, 3 rather small
posterodorsals near ;, 7;, and ¥, 1 distinct
ventral near % and sometimes 1 smaller
480
near 7;, usually 5 large apicals; hind tibia
with 7 anterodorsdals of irregular lengths
staggered in 2 rows, basal in series very
small, the second, fourth and sixth in a
more anterior row and slightly larger, with
6 posterodorsals, the third and fifth large,
near 7; and %, with 5 large apicals. Tarsus
with last segment slightly broadened, all
pulvilli enlarged and longer than Sth seg-
ment, fore tarsus slightly felted ventrally,
middle basitarsus with only a few ventral
spicules; lengths of tarsal segments from
the%base‘as: tarsus: 1,’0'52,,0:22.90575,0:13!
0.17 mm, pulvilli 0.26 mm; tarsus II, 0.65,
0.35, 0.22, 0.17, 0.13 mm, pulvilli 0.24
mm; tarsus III, 0.48, 0.43, 0.30, 0.17, 0.13
mm, pulvilli 0.20 mm.
Wings grayish with yellowish-brown
veins; anal margin fully rounded. Vein 1
reaching costa near middle of wing; vein 3
distinctly curved backward near tip; last
part of vein 4 broken at bend, broken ends
apparently slightly overlapping. Crossvein
% as long as terminal section of vein 5. Ca-
lypter, its setae and knob of halter whitish.
Abdomen metallic green dulled with gray
pollen; hairs mostly short, hairs on sides
and sternites longer, marginal setae 2 or 3
times as long as dorsal hairs. Genital cap-
sule dark brown, bearing 4 stout bristles
posteriorly.
Allotype female.—Length 4.0—4.5 mm;
wing 3.5—3.9 mm by 1.3 mm. Face slightly
higher than in male, with narrow clypeus
visible; antenna as in male. Fore and middle
coxae with setae less coarse anteriorly; fem-
ora without long setae or bristles below;
hind tibia with small anteroventrals; last
segments of tarsi not broader, pulvilli not
enlarged.
Holotype.—Male, JAMAICA: Falmouth,
bay shore, 1 March 1969, W. W. Wirth
(USNM).
Paratypes.—3 6, 3 @ (including alloty-
pe): same locality, date, collector, habitat as
holotype; | ¢, ANTIGUA: Dutchman Bay,
1 March 1969, W. W. Wirth (all USNM).
Etymology.—The species is named for
the collector, the late Willis W. Wirth of the
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Systematic Entomology Laboratory, Dip-
tera section of the U.S. Department of Ag-
riculture.
Diagnosis.—Asyndetus wirthi differs
from A. interruptus by the face of the male
being higher than wide, very gradually nar-
rowed from the front rather than having a
slight but abrupt narrowing at the level of
the antennae (Figs. 6, 7). The pedicel of the
antenna has the upper edge scarcely longer
than the lower edge, the first flagellomere
is scarcely as long as high with a very short
blunt point ventral to the arista, and the me-
soscutum has brownish pollen between the
rows of dorsocentrals. Asyndetus interrup-
tus has the pedicel of the antenna distinctly
longer above than below, the first flagello-
mere is swollen at the base with the lower
edge oblique to a short and sometimes
sharp point, and the mesoscutum is evenly
grayish or whitish pollinose above.
Discussion.—The distinction between
Asyndetus wirthi and Asyndetus interruptus
Loew was first made by Robinson (1975).
Unfortunately, at that time, A. wirthi was
thought to be A. interruptus, and specimens
of A. interruptus were described as a new
species, A. bredini. The confusion was the
direct result of inadequate knowledge of the
type specimen of A. interruptus, supposedly
deposited in the Poey collection in Havana,
Cuba. The type was said to be in a sealed
tray.
In 1990, this situation was resolved
through a correspondent in Cuba, Gabriel
Garcea G., who sent photographs of the
type specimen. Garcea reported that the
type was in the Gundlach collection of the
Ecology-Systematics Institute, Academy of
Sciences, in Havana, not in the Poey col-
lection. The specimen is a male, with the
genital capsule nearly hidden within the tip
of the abdomen. The antennae are now
missing from the type as depicted in the
photographs, and the colors were said to be
faded, but the head is present, and the fron-
tal and facial shapes can be seen. The face
is square, not higher than wide, the form
previously cited for A. bredini Robinson
VOLUME 99, NUMBER 3
481
Figs. 1-7.
4-6, A. wirthi. 4, Genital capsule. 5, Antenna. 6, Head showing face and front. 7, A. interruptus, head showing
face and front. 1, 4, 50 wm scale. 2-5, Upper 0.5 mm scale. 6, Lower 0.5 mm scale. 7, Drawn from photograph
of type specimen, seen from slightly above.
(1975). Asyndetus bredini is a synonym of
A. interruptus, and the material described as
A. interruptus in 1975 is described here as
a new species.
Both Asyndetus wirthi and A. interruptus
are found along seashores of the Gulf of
Mexico and Caribbean area, a habitat that
is common in the genus. The new species
is described strictly from Jamaica and An-
tigua, but it is presumed to occur more
Asyndetus. 1-3, A. archboldi. 1, Genital capsule. 2, Antenna. 3, Head showing face and front.
widely. A female, that may be this species,
has been seen from Quintana Roo, Mexico.
Asyndetus interruptus is known from Flor-
ida, Cuba and Dominica.
ACKNOWLEDGMENT
Gabriel Garcea G., at the Centro Oriental
de Ecosistemas y Bioversidad in Santiago
de Cuba, is thanked for the photographs of
482
the type specimen of Asyndetus interruptus
in Havana.
LITERATURE CITED
Abrahamson, W. G., A. E Johnson, J. N. Layne, and
P. A. Peroni. 1984. Vegetation of the Archbold
Biological Station, Florida: an example of the
southern Lake Wales Ridge. Florida Scientist 47:
209-250.
Cole, E R. 1969. The flies of western North America.
University of California Press, Berkeley. 693 pp.
Deyrup, M. 1990. Arthropod footprints in the sands
of time. Florida Entomologist 73: 529-538.
Deyrup, M. and T. Eisner. 1993. Last stand in the
sand. Natural History 102: 42—47.
1996. Description and natural history of a
new pygmy mole cricket from relict xeric uplands
of Florida (Orthoptera: Tridactylidae). Memoirs of
the Entomological Society of Washington 17: 59—
67.
Dyte, C. E. 1959. Some interesting habits of larval
Dolichopodidae (Diptera). Entomologists Monthly
Magazine 95: 139-143.
Myers, R. L. 1990. Scrub and high pine, pp. 150—
193. In Myers, R. L. and T. T. Ewel, eds. Ecosys-
tems of Florida. University of Central Florida
Press, Orlando. 765 pp.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Oldroyd, H. 1964. The natural history of flies. W. W.
Norton and Company, New York. 324 pp.
Robinson, H. 1964. A synopsis of the Dolichopodidae
(Diptera) of the southeastern United States and ad-
jacent regions. Miscellaneous Publications of the
Entomological Society of America 4: 103-192.
1975. Bredin-Archbold-Smithsonian biolog-
ical survey of Dominica, the family Dolichopod-
idae with some related Antillean and Panamanian
species (Diptera). Smithsonian Contributions to
Zoology 185: i-iv, 1-141.
Robinson, H. and J. R. Vockeroth. 1981. Dolichopod-
idae, pp. 625—639. Jn Manual of Nearctic Diptera.
1: i-vi, 1-674.
Smith, M. E. 1952. Immature stages of the marine fly,
Hypocharassus pruinosus Wh., with a review of
the biology of immature Dolichopodidae. Ameri-
can Midland Naturalist 48: 421—432.
Van Duzee, M. C. 1916. Table of males of the North
American species of the genus Asyndetus with de-
scriptions of six new species. Psyche 28: 88—94.
1919. Two new Asyndetus, with a table of
North American species. Entomological News 30:
248-250.
Williams, E X. 1938. Asyndetus carcinophilus Parent
(Diptera, Dolichopodidae). Proceedings of the Ha-
walian Entomological Society 10: 126-129.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 483-486
CRINODESSUS AMYAE, A NEW NEARCTIC GENUS AND SPECIES OF
PREDACEOUS DIVING BEETLE (COLEOPTERA: DYTISCIDAE:
HYDROPORINAE: BIDESSINI) FROM TEXAS, U.S.A.
KELLY B. MILLER
Department of Entomology, Colorado State University, Fort Collins, CO, 80523, U.S.A.
Abstract.—Crinodessus amyae n. gen. and n. sp. is described from southwestern Texas,
U.S.A. The genus is separable from all other genera of Bidessini by the combination of
1) cervical line posterior of and separated from posterior margin of the eye, 2) separation
of the genal line from the ventral margin of the eye broad, 3) apical segment of the
paramere elongate, 4) dense microreticulation on the ventral surface consisting of minute,
isodiametric cells, 5) relatively large size for members of the tribe, 6) distinctive habitus
with the pronotum cordate and the body elongate and depressed, and 7) anterior clypeal
margin extended anteriorly and prominently angulate. The similarity in habitus and other
characters between Crinodessus and Boongurrus Larson is hypothesized to be a result of
homoplasy due to similar behavior and common adaptation to a similar habitat. Liodessus
Guignot is tentatively proposed to be the sister genus of Crinodessus.
Key Words:
Since Bistré6m’s (1988) review of the
genera of the tribe Bidessini, two additional
genera have been described, Boongurrus
Larson (1994) from Queensland, Australia,
and Comaldessus Spangler and Barr (1995)
from Texas, U.S.A. Both are monotypic.
The following new species from south-
western Texas cannot be satisfactorily
placed into any existing genus of Bidessini.
Therefore, a new genus is described herein
to include it. The genus clearly belongs to
the tribe Bidessini based on the presence of
two-segmented parameres as defined by
Bistrom (1988).
The following description follows the
terminology and style of Larson (1994).
Measurements were taken with the aid of
an ocular micrometer in a Wild M3C dis-
secting microscope. Illustrations were made
using a drawing tube mounted to a Wild M3
dissecting microscope.
Taxonomy, predaceous diving beetle, Dytiscidae, new genus, Texas
Crinodessus Miller, new genus
Type species.—Crinodessus amyae, new
species.
Diagnosis.—The combination of the fol-
lowing characters is diagnostic for Crino-
dessus: 1) Separation of the transverse cer-
vical line and posterior margin of the eye;
2) separation of the genal line from the
margin of the eye broad; 3) apical segment
of the paramere elongate; 4) dense micro-
reticulation on the ventral surface of the
body consisting of minute, isodiametric
cells; 5) relatively large size for members
of the tribe; 6) distinctive habitus with cor-
date pronotum and elongate, depressed
body; and 7) prominently anteriorly extend-
ed and angulate anterior clypeal margin.
Description.—Small, length 2.58 mm;
body (Fig. 1) elongate; distinct discontinu-
ity between pronotum and elytron in dorsal
view; depressed. Color brown, slightly
484 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
dolene
3
4
1
Figs. 1-5. Crinodessus amyae. 1, Habitus. 2, Ventral surface of head, oblique view, showing genal line
(indicated by arrow). 3, Median lobe of aedeagus, dorsal view. 4, Median lobe of aedeagus, lateral view. 5,
Paramere, lateral view. Scale bar = 1 mm for Figs. 1 and 2 only.
darker on head and pronotum. Dorsal and
ventral surfaces moderately punctate, ven-
tral surfaces densely microreticulate.
Head: With clypeus prominently extend-
ed, forming angle with dorsal and ventral
surfaces; with a distinct transverse line (cer-
vical line) located between level of poste-
rior margin of compound eye and margin
of pronotum (Fig. 1), line relatively
straight; eyes relatively small, head width
between eyes/head width across eyes =
0.71; with a slightly depressed line near and
parallel to posteromedial margin of eye;
gena with distinct, curved line from pos-
terolateral margin of head to lateral edge of
base of maxilla (Fig. 2), region between eye
and line impressed, more strongly so in an-
terior portion; apical palpomere of each pal-
pus fusiform, apically bifid; antennomeres
2 and 3 subequal in length, antennomere 4,
0.6 length of 3, antennomeres 5—10 more
rounded and broader anteriorly than poste-
riorly.
Pronotum (Fig. 1): In dorsal aspect with
lateral margins strongly rounded, maximum
width anterior to middle, lateral margin
slightly sinuate basally with posterolateral
angle acute, posterior margin evenly curved
on each side of middle; lateral bead narrow
and of equal width throughout; disc with
sublateral, basal striae on each side of mid-
dle, stria extending about 0.6 length of pro-
notum measured along same general line as
Stria, stria slightly curved throughout
length; lateral margin curved downwards
on posterior angle. Scutellum not visible.
Elytron (Fig. 1): Without sutural stria;
basal stria present, originating slightly me-
dial to base of pronotal stria, about 0.5
length of pronotal stria, abruptly curved in
basal 0.25, thereafter straight and approxi-
mately parallel to elytral suture; disc strong-
ly flattened, without lateral grooves or ridg-
es; epipleuron broad and visible to base in
lateral aspect; not impressed basally, with-
out oblique, transverse carina; gradually
narrowed apically; evident to level of visi-
ble sternum 5. Hind wings present.
VOLUME 99, NUMBER 3
Prosternum: With anterolateral pore nar-
rowly separated from anterior margin; with
prominent, longitudinal, medial convexity
extending from anterior margin and contin-
uous with prosternal process; prominent se-
tae on sides of convexity. Prosternal pro-
cess in different plane than prosternum;
blade of process ovate, laterally sharply
margined, and setose; slightly convex in lat-
eral aspect; apex barely reaching antero-
medial lobe of metasternum, with minute
spine directed posteroventrally.
Metasternum: Metacoxae and metaster-
num combined length about 0.33 length of
body; metasternum with anteromedial pro-
cess rounded, not grooved, entire surface
moderately punctate without row of punc-
tures laterad to midline; metasternal wing
strongly arched and apically narrowed to
epipleuron; metacoxal lines very lightly im-
pressed, extending to metasternum, lines di-
vergent from near base, slightly less than
twice as long as distance between them at
narrowest point; metacoxal processes de-
pressed medially to level of abdominal ster-
num, each process with a minute, acute lobe
extending over base of trochanter.
Legs: Profemur with arched apicoventral
comb of yellow setae, its length about 0.21
length of profemur along ventral margin;
maximum width of protibia 0.30 of its
length, gradually widened from base to
apex; anterior surface of protibia smooth
and shiny with setae along margins; protar-
someres 1—3 broadly expanded, protarso-
mere 4 hidden in lobes of 3; metatrochanter
0.39 length of metafemur; metafemur with
maximum width near middle, narrowed
thereafter due to slight subapical convexity
along posterior margin, slightly widened
and rounded at apex; metafemur length 4.13
times greatest width; metatibia evenly wid-
ened apically, arcuate with point of greatest
curvature about 0.4 length from base, ven-
trally and mesally with natatorial setae;
metatarsus with natatorial setae, setae 1.38
times metatibial length.
Genitalia: Median lobe of aedeagus
(Figs. 3, 4) simple, evenly arcuate in lateral
485
aspect; lateral margins slightly divergent to
rounded apex in dorsal aspect; paramere
(Fig. 5) 2-segmented, apical segment about
0.79 length of basal segment, with a min-
ute, hook-like apical projection. Female and
larva unknown.
Etymology.—The masculine generic
name is derived from the Greek word crino,
“‘separate,”’ referring to the separation be-
tween the cervical line and the posterior
margin of the eye, and dessus, the suffix of
the type genus of the tribe, Bidessus Sharp.
Discussion.—Crinodessus shares several
characters with Boongurrus Larson includ-
ing the elongate and flattened shape of the
body, the cordate pronotum and the broad
separation of the genal line from the margin
of the eye. However, Crinodessus differs
from Boongurrus in having an angulate an-
terior margin of the clypeus, an elongate
apical segment of the paramere, and sepa-
ration of the cervical line and the posterior
margin of the eye. In addition, Boongurrus
lacks microreticulation on the ventral sur-
face, is more densely punctate and is small-
er. It seems likely that the similarity be-
tween these genera is a result of homoplasy
rather than close relationship. According to
Larson (1994), small eyes, cordate prono-
tum, and elongate flattened shape are char-
acteristic of species which live in substrate
or move through interstices, the habitat of
Boongurrus (Larson 1994) and the pre-
sumed habitat and behavior of Crinodessus.
The broad separation of the genal line from
the margin of the eye and separation of the
cervical line from the eye in Crinodessus
may have resulted from reduction in size of
the eye and, partly, from elongation of the
body. If Crinodessus and Boongurrus are
closely related, it would be an unusual re-
lationship given the broadly disjunct distri-
bution of the two genera.
The simple shape of the genitalia along
with the presence of a transverse cervical
line and lack of sutural line on the elytra in
Crinodessus indicate a possible sister-group
relationship with Liodessus Guignot. Cri-
nodessus keys to Liodessus in Young’s
486
(1967) and Bistr6m’s (1988) keys to genera
of Bidessini. However, Crinodessus differs
from Liodessus in all of the above diagnos-
tic characters except elongate apical seg-
ment of the paramere, a presumably ple-
siomorphic character (Larson 1994). The
relationship between these two genera is,
therefore, not well founded. However, Cri-
nodessus shares no obvious affinities with
any other genus.
Crinodessus amyae Miller, new species
(Figs. 1-5)
Holotype.—Male: labelled ““TEXAS:
Presidio County, 12 mi NE Ruidosa, Pinto
Canyon, 23 October 1985, Rawlins & Da-
vidson, legs /HOLOTYPE Crinodessus
amyae K. B. Miller 1997 [red label].’’ De-
posited in The Carnegie Museum of Natural
History. Specimen dissected with genitalia
in microvial of glycerin mounted on pin.
According to R. Davidson (in litt.), the
specimen was collected from a clear desert
stream which flows over clean sand and
gravel in the Chinati Mountains of south-
western Texas. The stream is about a meter
wide with a few deeper pools.
Description.—Measurements: Length
2.58 mm; maximum width 1.19 mm; 2.17
times longer than wide, pronotum 0.85
times maximum width of body.
Color: Head brown, slightly darker brown
anteromesad of each eye, slightly lighter on
surface of frons and along margin of each
eye; 4 basal antennal segments yellow,
brown thereafter; palpi yellow-brown; pro-
notum brown, darker medially and gradu-
ally more pale laterally; elytra slightly light-
er brown than head and pronotum, without
maculations; ventral surface uniformly yel-
low-brown.
Sculpture: Head shiny, lightly shagree-
ned and finely and evenly punctate; no
punctures posterior to transverse line. Pro-
notum shiny, anterolateral surface slightly
shagreened; moderately and evenly punc-
tate; punctures larger than on head and sep-
arated by about 1.5 times puncture diameter
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
except less dense mediad to apical half of
each stria; slightly longitudinally rugulose
mediad to base of striae. Elytron shiny and
smooth; punctures similar in size and dis-
tribution to those of pronotum; punctation
evident immediately adjacent to suture;
slightly rugulose mediad to bases of striae.
Epipleuron lightly shagreened and moder-
ately punctate; punctures separated by 1.5—
2.0 times puncture diameter and with short,
decumbent setae. Metasternum, metacoxae
and abdominal sterna with distinct micro-
reticulation of minute, isodiametric cells
covering entire surface; microreticulation
less conspicuous on abdominal sterna.
Etymology.—I take pleasure in naming
this species in honor of my friend and wife,
Amy, who has been a source of great sup-
port during my studies of Dytiscidae.
ACKNOWLEDGMENTS
I thank B. C. Kondratieff for reviewing
the manuscript and for his encouragement
and support. S. Fitzgerald, P. Opler, R. E.
Roughley and an anonymous reviewer also
provided valuable comments on the manu-
script. Thanks to R. Davidson and D. Pol-
lock for the opportunity to examine the col-
lection of Dytiscidae at The Carnegie Mu-
seum of Natural History, Pittsburgh, PA.,
which led to the discovery of the specimen.
Davidson also provided biological infor-
mation. Special thanks to A. Humphrey and
P. Shephard of The Nature Conservancy for
the use of their facilities during the writing
of this manuscript.
LITERATURE CITED
Bistr6m, O. 1988. Generic review of the Bidessini
(Coleoptera, Dytiscidae). Acta Zoologica Fennica
184: 1-41.
Larson, D. J. 1994. Boongurrus rivulus, a new genus
and species of water beetle (Coleoptera: Dytisci-
dae: Bidessini) from northern Queensland, Austra-
lia. Journal of the Australian Entomological So-
ciety 33: 217-221.
Spangler, P. J. and C. B. Barr. 1995. A new genus and
species of stygobiontic dytiscid beetle, Comaldes-
sus stygius (Coleoptera: Dytiscidae: Bidessini),
from Comal Springs, Texas. Insecta Mundi 9:
301-308.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 487-489
GALL MIDGE FORMS GALLS ON FRUIT FLY GALLS
(DIPTERA: CECIDOMYIIDAE, TEPHRITIDAE)
DAviIpD H. HEADRICK AND RICHARD D. GOEDEN
Department of Entomology, University of California, Riverside, CA 92521, U.S.A.
Abstract.—A cecidomyliid gall midge, Rhopalomyia bigeloviae (Cockerell), was reared
from galls formed on galls of the tephritid, Aciurina trixa Curran, on Chrysothamnus
nauseosus (Pallas) Britton in southern California. Only one of the two A. trixa gall types
in southern California hosted the gall midge. Rhopalomyia bigeloviae galls were originally
discovered in Colorado on the large cottony galls of A. bigeloviae, a species closely related
to A. trixa, that forms galls on C. nauseosus outside of California. The midge galls are
similar to the host tephritid galls externally, i.e. either smooth or cottony. The number of
midge galls per tephritid gall varies, but they can, in some cases, cover the entire surface
of the tephritid gall. The gall midge is probably bivoltine, and the tephritid host is uni-
voltine. Gall midge galls first become visible in early spring (March). Adults emerge later
in the spring. The fate of these adults remains unknown, but females may oviposit into
other parts of the host plant to form a second generation of cecidomyiid galls not depen-
dent on the presence of a tephritid gall.
Key Words:
palomyia, bigeloviae
Aciurina trixa Curran forms axillary bud
galls on branches of its only known host
plant, Chrysothamnus nauseosus (Pallen)
Britton, in western United States; but, the
morphology of its gall varies strikingly
within its range. In Idaho, at least three gall
morphs of A. trixa were reported by Wang-
berg (1981); in southern California, we rec-
ognize two gall morphs for A. trixa on C.
nauseosus (Headrick et al. 1997).
On one of the two gall forms in southern
California (Fig. 1A), we observed that its
exterior sometimes was covered by small
nodules at different densities among differ-
ent locations. Upon dissection, each of
these nodules was found to contain a small
cecidomyiid identified for us as Rhopalo-
myia bigeloviae (Cockerell) by Raymond J.
Gagné (Systematic Entomology Labora-
tory, Agricultural Research Service,
Gall formers, gall midge, Cecidomyiidae, Tephritidae, Aciurina, trixa, Rho-
USDA). This species was originally de-
scribed from a large woolly gall up to 12
mm long, typical of that made by another
species of Aciurina (Gagné 1986). Dodson
and George (1986) and Headrick et al.
(1997) have clarified the status of Aciurina
species on C. nauseosus, and this woolly
gall is now attributed to the tephritid, A.
bigeloviae (Cockerell). We have examined
such woolly galls from various collections
(Washington State University, Pullman;
University of Idaho, Moscow; G. Dodson,
personal collection) and they do, indeed,
bear R. bigeloviae galls. None of the other
gall types, as described by Wangberg
(1981), examined from museum collections
throughout western U.S., including the
smaller resinous gall of A. trixa found in
California hosted this gall midge (Headrick
et al. 1997).
488
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ioe ie
In southern California, the midge galls
become visible in early spring (March) and
grow very rapidly (Fig. 1B), as do other
Rhopalomyia spp. galls (Gagné 1989). The
gall in saggital section is layered exactly
like the tissues comprising the parent A.
trixa gall, including the smooth, waxy, out-
er layer. On the cottony galls of A. bigelov-
iae, the midge galls also bear the thick to-
mentum of the host fruit fly gall, thus, the
(A) The fully formed gall of Aciurina trixa on Chrysothamnus nauseosus; (B) detail of the cecido-
myiid, Rhopalomyia bigeloviae, galls on an A. trixa gall; (C) saggital section of A. trixa and R. bigeloviae galls;
(D) detail of the R. bigeloviae gall with cecidomyiid larva within; (E) the exposed pupa of R. bigeloviae; (F)
an adult female of R. bigeloviae.
midge gall takes on the surface character-
istics of the host gall. In southern Califor-
nia, the midge larvae feed within their gall
locules as the tephritid larva continues to
develop in its own, much larger, separate
locule (Fig. 1 C, D). Midge development
proceeds rapidly and both the tephritid and
midge co-occur as late-instar larvae, and
pupate at about the same time. The midge
pupa forms inside its gall locule, but the
VOLUME 99, NUMBER 3
waxy apex of its gall sometimes splits, part-
ly exposing the pupa within (Fig. E). The
gall midge adult emerges in April (Fig. FP),
but the fate of this generation remains un-
known. We know that A. frixa is univoltine
on C. nauseosus, with the gall dying after
emergence of the fruit fly adult. Thus, the
current season’s tephritid galls are no longer
suitable for oviposition by the midge. The
options for the adult female gall midge fol-
lowing emergence include, but are not re-
stricted to, oviposition into the primordial
tephritid gall tissues or the axillary bud
galls of the host plant or oviposition on an-
other part of the host plant, thus cycling
between alternate gall types. The latter ap-
pears likely as this has been reported for
other species in this genus (Gagné 1989).
There is as yet no indication that the ceci-
domyiid affects the growth and develop-
ment of the tephritid and the relationship
thus appears unequal, with the cecidomyiid
dependent on the tephritid during this part
of its life cycle.
This is the first known description and
illustration of a cecidomyiid gall being
formed upon a tephritid gall. Tephritids are
known to be inquilines of cecidomyiid galls
(Jones et al. 1983), but this is the most in-
timate association between gall-forming
flies of these two families known to us.
Some workers may not have previously
recognized this phenomenon and interpret-
ed the growths on tephritid galls as devel-
oping leaves (Fernandez and Price 1994).
This does happen, but closer inspection
489
may otherwise reveal not a budding leaf,
but rather a developing cecidomyiid gall!
Dedication.—We would like to dedicate
this paper to Raymond J. Gagné in honor
of his recent retirement from the Systematic
Entomology Laboratory, U.S. Department
of Agriculture.
LITERATURE CITED
Fernandez, G. W. and P. W. Price. 1994. Life history,
courtship, and mating behavior of the gall-forming
Aciurina trixa (Diptera: Tephritidae) on Chryso-
thamnus nauseosus hololeucus (Asteraceae). Pro-
ceedings of the Entomological Society of Wash-
ington 96: 301—307.
Gagné, Raymond. J. 1989. The plant-feeding gall
midges of North America. Comstock Publishing
Associates, Cornell University Press, Ithaca and
London. 356 pp.
Goeden, R. D., D. H. Headrick, and J. A. Teerink.
1995. Life history and description of immature
stages of Valentibulla californica (Coquillett)
(Diptera: Tephritidae) on Chrysothamnus nauseo-
sus (Pallas) Britton in southern California. Pro-
ceedings of the Entomological Society of Wash-
ington 97: 548-560.
Headrick, D. H., R. D. Goeden, and J. A. Teerink.
1997. Taxonomy of Aciurina trixa Curran and its
life history on Chrysothamnus nauseosus in
southern California; with notes on A. bigeloviae
(Cockerell). Proceedings of the Entomological So-
ciety of Washington 99: 415—428.
Jones, R. G., R. J. Gagné, and W. FE Barr. 1983. A
systematic and biological study of the gall midges
(Cecidomylidae) of Artemisia tridentata Nuttall
(Compositae) in Idaho. Contributions of the
American Entomological Institute 21(2): 1—79.
Wangberg, J. K. 1981. Gall-forming habits of Aciuri-
na species (Diptera: Tephritidae) on Rabbitbrush
(Compositae: Chrysothamnus spp.) in Idaho. Jour-
nal of the Kansas Entomological Society 54: 711—
IBY.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 490-493
FOOD LIMITATION REDUCES BODY LENGTH IN MANTID NYMPHS,
TENODERA SINENSIS SAUSSURE (MANTODEA: MANTIDAE):
IMPLICATIONS FOR FITNESS
KATHLEEN DUSSE AND L. E. HuURD
Department of Biology, Washington & Lee University, Lexington, VA 24450, U.S.A.;
LEH Correspondence author.
Abstract.—Growth rate and body size have been linked to fitness in the mantid, Ten-
odera sinensis Saussure (Mantodea: Mantidae). We asked how early in the life cycle food
level could affect these two parameters. Two laboratory cohorts were offered prey at either
high or low density during first and second stadia. These nymphs exhibited significant
differences in predation rate, growth rate, and body size, but not in gross growth effi-
ciency. Well-fed nymphs achieved larger body size in a shorter time than poorly fed ones
during both their first and second stadia. Because body size of adult females determines
maximum fecundity, this response suggests that food level during the early life history
of this species can directly affect fitness. Flexibility in body size and rate of development
may play an important role in determining distribution and regional persistence of this
species in temperate habitats.
Key Words:
sinensis
For generalist predators such as praying
mantids, limited prey availability in nature
is common, and can reduce growth, surviv-
al, and reproduction (Matsura 1981, Mat-
sura and Marooka 1983, Hurd and Eisen-
berg 1984, Hurd et al. 1995). The impact
of food limitation on growth and develop-
ment would seem particularly acute for
temperate zone mantids, that are faced with
a limited growing season in which to com-
plete their life cycles before freezing weath-
er kills the adults.
Tenodera sinensis (Saussure) was intro-
duced into this country in 1896 near Phil-
adelphia, PA (Laurent 1898). It has since
become a widespread and abundant mantid
in the eastern United States, with a range
extending from southern New England
through the Carolinas (Rooney et al. 1996).
This large-bodied species hatches from con-
Body size, fitness, food limitation, growth, Mantidae, Mantodea, Tenodera
tagiously dispersed oothecae early in the
spring, leading to dense concentrations of
hatchlings when prey levels are low (Eisen-
berg and Hurd 1990). Low prey availability
reduces both rate of development (Hurd and
Rathet 1986) and survival (Hurd and Eisen-
berg 1984) during the early stadia. This is
a semelparous, univoltine species which
overwinters in the egg stage. Thus, reduced
rate of development can retard maturation
so that females do not have time to com-
plete seven nymphal stadia, undergo oogen-
esis, and oviposit before killing frost (Hurd
et al. 1995).
One consequence of early food limitation
is reduced imago body length, and body
length of adult females has been implicated
as the limiting factor in egg production. Ei-
senberg et al. (1981) found that adult fe-
males with longer mean body length gained
VOLUME 99, NUMBER 3
more mass in the field during oogenesis
than those that had lower mean body length
at a different site. Mass gain at this time of
the life cycle is directly related to number
of eggs contained in oothecae (Eisenberg et
al. 1981, Hurd 1989). Thus, feeding con-
dition of juvenile mantids can affect fitness
(egg production) through body length. We
asked how early in the life cycle the effect
of food availability on components of fit-
ness might be evident, by raising hatchling
Tenodera sinensis in the laboratory at two
prey abundance levels and recording body
lengths at ecdysis and time spent in differ-
ent stadia.
MATERIALS AND METHODS
Thirty-nine oothecae of Tenodera sinen-
sis were collected early in the spring of
1996 from old fields in Rockbridge Co., Va.
Four experimental groups of 20 nymphs
each were established from individuals that
hatched the same day. These groups were
fed apterous Drosophila melanogaster
(Meigen) at two levels and killed by freez-
ing to measure and weigh after either the
first or second molt.
Groups A and B were killed after the first
molt. Nymphs in group A were fed four
flies per day; those in group B were fed one
fly per day. Groups C and D were killed
after the second molt. Nymphs in group C
were fed four flies per day; those in group
D were fed one fly per day.
All nymphs were housed individually in
130-ml glass vials having tight lids fitted
with fine-mesh brass screening for ventila-
tion, and a hole through which a cotton
swab, moistened daily with distilled water,
was placed (Hurd and Rathet 1986). All vi-
als were kept in a growth chamber set at
23°C and on a 16:8 (L:D) light cycle.
Body length, from front of head to tip of
abdomen, was measured to the nearest 0.1
mm for all killed nymphs after thawing.
Dry weight body mass was measured to the
nearest 0.1 mg after drying in an oven at
50°C for 24 h. Number of days in a stadium
was recorded for each nymph.
49]
Because feeding rate can affect the effi-
ciency with which food is processed by
these predators (Hurd 1991), we calculated
gross growth efficiency (GGE) for mantid
nymphs in the first stadium, for groups A
and B:
GGE = [(mass gained)
+(mass of food consumed)]
xX 100%
where mass gained was individual mass of
a nymph at ecdysis minus an established
mean dry mass of 17 nymphs from the orig-
inal cohort that had been killed, dried, and
weighed upon hatching. Mass of food con-
sumed was calculated by multiplying the
number of flies eaten during a stadium by
the mean dry weight mass measured from
50 flies taken from culture bottles at the be-
ginning of the experiment. GGE could not
be calculated for second stadium nymphs in
C and D, because we had no baseline for
beginning second instars from these two co-
horts.
RESULTS
Mantids in both first and second stadia
ate more, developed faster, and attained
greater mean body length when fed four
flies per day than when fed only one per
day (Table 1). Group A and C nymphs fed
at three to four times the rate of those in
groups B and D, respectively.
Group B nymphs took about 50% longer
to complete the first stadium than those in
group A. The difference in rate of devel-
opment was even more pronounced during
second stadium: group D took nearly three
times as long to complete the stadium as
group C. In contrast, well-fed nymphs com-
pleted both first (A) and second (C) stadia
in about the same time.
The percent gain in body length of well-
fed nymphs compared to poorly fed
nymphs increased from first to second sta-
dium. Well-fed group A nymphs in first sta-
dium attained a mean of 11% greater body
length than poorly-fed group B individuals.
For nymphs completing second stadium,
492 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Mean (+SE) predation rate (flies/day eaten), development time (days in stadium), and body length,
for first and second stadium Tenodera sinensis nymphs, mean (+SE) percent gross growth efficiency (GGE) for
first instars, and results of 2-tailed rt tests for differences between pairs of groups. Calculation of GGE for first
stadium nymphs explained in text. GGE was not significantly different between A and B after table-wide P
adjustment for multiple comparisons (Rice 1989).
Group Flies Eaten/Day Days in Stadium Body Length (mm) %GGE
A 2.6 (+£0.06) ts) ((22(0)5113))) 152275 (=0F1)) 30.8 (£0.99)
B 0.9 (£0.01) WAI (Ess) NEI57/ (22h I)) 27.4 (+£1.24)
A v. B: tig ~29.83 ~9.15 ~9.02 ~2.14
P= <0.001 <0.001 <0.001 0.04 (NS)
€ BESnGs0!05) 8.2 (+£0.14) 20.5 (0:21)
D LOKG=OOD) 22.8 (+£0.74) 16.8 (£0.15)
(Ee IDE fas 54.67 = IID LIS 13.88
P= <0.001 <0.001 <0.001
mean body length of group C nymphs was
22% greater than of those in group D. In
fact, poorly fed group C nymphs complet-
ing second stadium (D) were hardly larger
than well-fed group A nymphs completing
first stadium.
Feeding level did not significantly affect
GGE, though nymphs in group A were
slightly more efficient on average than
those in B (Table 1).
DISCUSSION
The benefit from increased feeding op-
portunity in 7. sinensis nymphs occurred
immediately after eclosion, in terms of both
size and rate of development. The advan-
tage in body size during first stadium in-
creased during second stadium, indicating
that the increased body size of well-fed first
instars accumulates as individuals grow, at
least through the second stadium. This
could explain differences in adult body
length (7—10 cm) in a given field population
(personal observations). To the extent that
body length determines maximum egg pro-
duction, early feeding clearly can have a
significant impact on fitness of females in
this species.
Sex cannot be determined in T. sinensis
until the third stadium, but sex ratio of
hatchlings is 50:50 (Moran and Hurd 1994).
However, although this means that half of
our experimental animals were male, the ef-
fect of body size on male fitness has not yet
been demonstrated.
The impact of feeding on development
rate was also marked. For well-fed nymphs,
the time to complete first and second stadia
was about the same (8 days), although
nymphs fed a single fly per day took nearly
twice as long to complete second stadium
as compared to first. Thus, well-fed nymphs
took approximately 16 days (total) to com-
plete the first two stadia; poorly fed
nymphs, on the other hand, took more than
30 days to do the same thing. Our data sug-
gest that well-fed nymphs could nearly
complete four stadia in 30 days. This man-
tid is an ambush predator, and its nymphs
do not move about very much in the field
(Rathet and Hurd 1983). Thus, we feel it is
unlikely that our nymphs expended signif-
icantly less energy in the vials (i.e., have
more energy to grow) than they would have
in the field.
Hurd (1991) found that %GGE varied
with predation rate in first stadium T. si-
nensis nymphs: from 34% for <0.4 flies per
day to 57% for 0.8 flies per day, reduced to
about 40% for 1.2 flies per day. However,
in the present experiment mean %GGE did
not significantly differ between groups that
ate 0.9 flies/day and those that ate 2.6 flies
per day. In fact, %GGE was lower in this
experiment than for any group in the earlier
study. The design of this experiment was
VOLUME 99, NUMBER 3
similar but there may have been a proce-
dural difference, a genetic difference be-
tween these mantid nymphs and those used
in the earlier study, or both that caused the
disparity in results.
The degree of phenotypic flexibility of
body size and rate of development exhib-
ited by this species from the earliest stage
of development has important implications
for its distribution and persistence. Clearly,
its latitudinal distribution will be limited to
habitats of sufficient season length and
feeding opportunity. However, the present
contiguous distribution of this species in the
eastern U.S. (approximately from southern
Pennsylvania to northern Georgia) indicates
a wide tolerance with regard at least to
length of growing season. Because its body
size is flexible, this species should be able
to complete its life cycle under a variety of
food levels. Thus, though early feeding ex-
perience can affect fitness in this species,
flexibility in adult body length may partial-
ly mitigate the predicted shrinkage of re-
gional distribution under global warming
scenarios (Rooney et al. 1996).
The salient features of the life history of
this mantid are shared by many other pre-
dacious arthropods inhabiting temperate
seasonal environments. Our findings that
adult fitness can be influenced by food
availability immediately upon egg hatch
may therefore be more general than for T.
sinensis. However, we know of no compre-
hensive studies linking food limitation,
growth rates, body size, and fitness in other
such predators.
ACKNOWLEDGMENTS
The first author was supported by an R.
E. Lee Research Fellowship from Washing-
ton & Lee University.
LITERATURE CITED
Eisenberg, R. M. and L. E. Hurd. 1990. Egg disper-
sion in two species of praying mantids (Mantodea:
493
Mantidae). Proceedings of the Entomological So-
ciety of Washington 92: 808-810.
Eisenberg, R. M., L. E. Hurd, and J. A. Bartley. 1981.
Ecological consequences of food limitation for
adult mantids (Tenodera aridifolia sinensis Saus-
sure). American Midland 106: 209—
218.
Hurd, L. E. 1989. The importance of late season flow-
ers to the fitness of an insect predator, Tenodera
sinensis Saussure (Orthoptera: Mantidae), in an
old field community. Entomologist (London) 108:
223-228.
Hurd, L. E. 1991. Growth efficiency in juvenile man-
tids: absence of selection for optimization in a
food-limited environment (Orthoptera: Mantidae).
Proceedings of the Entomological Society of
Washington 93: 748-750.
Hurd, L. E. and R. M. Eisenberg. 1984. Experimental
density manipulations of the predator Tenodera si-
nensis (Orthoptera: Mantidae) in an old-field com-
munity. I. Mortality, development and dispersal of
juvenile mantids. Journal of Animal Ecology 53:
269-281.
Hurd, L. E., R. M. Eisenberg, M. D. Moran, T. P. Roo-
ney, W. J. Gangloff, and V. M. Case. 1995. Time,
temperature, and food as determinants of popula-
tion persistence in the temperate mantid Tenodera
sinensis (Mantodea: Mantidae). Environmental
Entomology 24: 348-353.
Hurd, L. E. and I. H. Rathet. 1986. Functional re-
sponse and success in juvenile mantids. Ecology
67: 163-167.
Laurent, R. 1898. A species of Orthoptera. Entomo-
logical News 9: 144-145.
Matsura, T. 1981. Responses to starvation in a mantis,
Paratenodera angustipennis (S.). Oecologia 50:
291-295.
Matsura, T. and K. Marooka. 1983. Influence of prey
density on a mantis, Paratenodera angustipennis
(S.). Oecologia 56: 306-312.
Moran, M. D. and L. E. Hurd. 1994. Experimentally
determined male-biased sex ratio in a praying
mantid. American Midland Naturalist 132: 205—
208.
Rathet, I. H. and L. E. Hurd. 1983. Ecological rela-
tionships of three co-occurring mantids, Tenodera
sinensis (Saussure), T. angustipennis (Saussure),
and Mantis religiosa (Linnaeus). American Mid-
land Naturalist 110: 240-248.
Rice, W. R. 1989. Analyzing tables of statistical tests.
Evolution 43: 223-225.
Rooney, T. P, A. T. Smith, and L. E. Hurd. 1996.
Global warming and the regional persistence of a
temperate-zone insect (Tenodera sinensis). Amer-
ican Midland Naturalist 136: 84—93.
Naturalist
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 494-501
NEW SPECIES OF SOUTHWESTERN NEARCTIC MICROCADDISFLIES
(TRICHOPTERA: HYDROPTILIDAE)
STEPHEN R. MOULTON II AND STEVEN C. HARRIS
(SRM) U.S. Geological Survey, National Water Quality Laboratory, Biological Unit,
5293 Ward Road, MS 407, Arvada, CO 80002, U.S.A.; (SCH) Department of Biology,
Clarion University, Clarion, PA 16214, U.S.A.
Abstract.—Four new species of Hydroptilidae are described and illustrated. Hydroptila
abbotti n. sp. and Ochrotrichia boquillas n. sp. are known from the Austroriparian and
Chihuahuan biotic provinces of Texas, respectively. Oxyethira garifosa n. sp. and Oxy-
ethira desadorna n. sp. are described from northern Mexico. The new species are diag-
nosed with closely related congeners.
Key Words:
Southwestern Nearctic caddisflies are
poorly known. With recent concentrated
survey efforts in Texas by Moulton and
Stewart (in press) and sporadic collections
in northern Mexico, species new to science
continue to be discovered. This is especially
apparent for the family Hydroptilidae. This
paper describes four new species of micro-
caddisflies, one each in the genera Hydrop-
tila and Ochrotrichia, and two in the genus
Oxyethira. In North America Hydroptila is
the most diverse hydroptilid genus with 110
species; Ochrotrichia and Oxyethira con-
tain 72 and 40 species, respectively (Morse
1993). This family includes the smallest
species of caddisflies, ranging from 1.5 to
5 mm. All of the material examined in this
study was collected with an ultraviolet light
trap.
Length is measured from the top of the
head to the tip of the forewing. Holotypes
are deposited in the National Museum of
Natural History, Smithsonian Institution
(NMNH). Paratypes are deposited in the
NMNH, the Academy of Natural Sciences
of Philadelphia (ANSP), the Illinois Natural
History Survey (INHS), and the collections
of the authors (SRM, SCH).
Trichoptera, Hydroptilidae, microcaddisflies, Texas, Mexico, new species
Hydroptila abbotti Moulton and Harris,
new species
(Fig. 1)
Hydroptila n. sp.: Moulton and Stewart, in
press.
Description.—Length 2.0 mm. Brown in
alcohol. 25 antennal segments. Segment VII
with short ventromesal process, apex
rounded in lateral view, triangular in ventral
view. In lateral view, segment VIII some-
what triangular; in dorsal view, posterior
margin incised. Segment IX short, retracted
within segment VIII; in lateral view, ante-
rior margin broadly rounded, narrowing
posterad. Inferior appendages long, reach-
ing to upturned portion of X; in lateral
view, angled ventrad, ventral margin cren-
ulate with apex pointed and heavily scler-
otized; in ventral view, mesal margin ridge-
like bearing four dark, stout points apically,
several smaller dark points basally, broadest
basally and narrowing towards shoulder-
like apex. Segment X lightly sclerotized,
approximately three times longer than wide;
in dorsal view, widest in middle, apical
one-third deeply bifurcate, tips of bifurca-
VOLUME 99, NUMBER 3 495
Fig. 1. Hydroptila abbotti, male genitalia. A, Left lateral view. B, Dorsal view. C, Ventral view. D, Phallus.
496
tions darkly pigmented and serrate; in lat-
eral view, concave in middle, apices acute
and angled dorsad. Phallus needle-like, ba-
sal one-quarter broadest; spiraled paramere
making one revolution around shaft at mid-
length.
Immature stages and female-—Unknown.
Holotype male.—U.S.A.; TEXAS, An-
derson Co., Skeet Branch, Engeling Wild-
life Management Area, 3.2 km W Black-
foot, 12 June 1994, J. Abbott, J. Chirhart,
M. Pasanante (NMNH).
Etymology.—Named for John C. Abbott,
collector of the holotype.
Diagnosis.—This species belongs to the
H. waubesiana group of Marshall (1979)
and is most closely related to the recently
described H. homochitta Harris and Sykora
(Harris and Sykora 1996). Hydroptila ab-
botti is distinguished from the latter by the
upturned, acutely tapered apex of tergum X
in lateral view (club shaped in H. homo-
chitta); the narrower base of tergum X in
dorsal view, and the more developed api-
colateral shoulder of each inferior append-
age in ventral view. Additionally, unlike H.
homochitta, H. abbotti does not possess a
pair of long, thin intermediate appendages
beneath tergum X.
Distribution.—Known only from the
type locality in the east Texas Gulf Coastal
Plain. This location is a first order, spring-
fed, sand-bottomed stream. Hydroptila ab-
botti was collected along with H. ouachita
Holzenthal and Kelley, a species previously
known only from Schoolhouse Spring,
Jackson Parish, Louisiana (Holzenthal and
Kelley 1983), approximately 480 km east
of the Texas collection locality. Both of
these locations have similar habitat char-
acteristics.
Ochrotrichia boquillas Moulton and
Harris, new species
(Fig. 2)
Ochrotrichia n. sp.: Moulton and Stewart,
in press.
Description.—Length 2.5 mm. Brown in
alcohol. 29 antennal segments. Segment VII
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
with short, acute ventromesal process. Seg-
ment VIII rectangular in dorsal and lateral
views. Segment IX in lateral view with pos-
terior margin broadly rounded; anteroven-
tral margin produced slightly into segment
VIII. In lateral view, inferior appendages
lobate, curving dorsad; in ventral view, bas-
es broad, ventromesal surfaces shoulder-
like in middle, diverging towards apices,
each lined with comb-like row of erect,
stout setae; apices with row of longer, hair-
like setae. In dorsal view, segment X spat-
ulate, rounded on apex; wedge-shaped in
lateral view; basodorsal area bearing two
sclerotized processes, left process V-sha-
ped, right process ventrolaterally hooked.
Phallobase flared, tapering towards middle;
apical one-half nearly parallel sided; apex
with prominent ejaculatory duct emerging
from middle, encircled preapically by acute,
sclerotized process.
Immature stages and female-—Unknown.
Holotype male.—U.S.A.; TEXAS, Brew-
ster Co., Glenn Spring, Big Bend National
Park, 18 April 1993, R. Garano (NMNBH).
Paratypes.—Same as holotype, but Rio
Grande Village Campground, 2 April 1993,
J. Gelhaus and D. Koenig, 4 d (ANSP); Val
Verde Co., Dolan Creek at Penstemon Cliff
Springs above confluence Devils River,
27-28 July 1995, G. Easley, 1 6 (NMNH).
Etymology.—Named for Boquillas Can-
yon, through which the Rio Grande flows
near the Big Bend area of Texas and Mex-
ico.
Diagnosis.—Ochrotrichia boquillas is a
member of the O. xena group (Flint 1972)
and is most closely related to O. flagellata
Flint and O. pectinata Flint. Ochrotrichia
boquillas differs from these species by the
lobate structure of the inferior appendages
in lateral view and the configuration of spi-
nous processes on tergum X.
Distribution Known only from the lo-
calities of the type material.
Oxyethira garifosa Moulton and Harris,
new species
(Fig. 3)
Description.—Length 2.8 mm. Brown in
alcohol. 40 antennal segments. Segment VII
VOLUME 99, NUMBER 3 497
Fig. 2. Ochrotrichia boquillas, male genitalia. A, Left lateral view. B, Dorsal view. C, Ventral view. D,
Phallus.
498 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Oxyethira garifosa, male genitalia. A, Left lateral view. B, Dorsal view. C, Ventral view. D, Phallus.
VOLUME 99, NUMBER 3
with short, acute ventromesal process. Seg-
ment VIII with lateral margins produced
posterad, apex truncate, slightly scalloped;
dorsal and ventral margins broadly emar-
ginate. Segment IX complete dorsally and
concealed within segments VII and VIII,
anterior one-half extending to posterior
margin of segment VI, posterior margin sin-
uate with broad lobe ventrally bearing sev-
eral stout setae. Tergum X membranous in
lateral view; rectangular in dorsal view. In-
ferior appendages fused: in lateral view,
dorsal margin twice as long as ventral mar-
gin, concave on posterior margin; in ventral
view, appearing as four lobate processes.
Subgenital plate narrow in lateral view, dis-
tally with acute ventral tooth; narrowing
mesally in ventral view, sclerotized along
posterior margin. Bilobed processes widely
separated in ventral view. Phallobase tubu-
lar, tapering in middle; apical one-half di-
vided into two sclerotized processes, short-
er process sinuate, apically acute, longer
process broadest basally, constricted and
twisted in middle, recurving to an acute
apex.
Immature stages and female.—Unknown.
Holotype male.-—MEXICO; TAMAU-
LIPAS, Municipio de Ciudad Victoria, Ar-
royo los Troncones, Ejido La Libertad, ca.
10 km NW Victoria, 14 May 1989, S. Har-
ris, A. Contreras, and A. Moreno (NMNH).
Etymology.—Spanish; hook-like, refer-
ring to the distinctive phallic structure.
Diagnosis.—This species fits within the
Damphitrichia subgenus in the pallida
group as established by Kelley (1984)
based upon the complete dorsum of seg-
ment IX and the absence of a spiraled par-
amere on the phallus. The new species is
most similar to O. arizona Ross and O. ver-
na Ross in overall appearance. From the
former, O. garifosa is distinguished by the
absence of dorsolateral processes on seg-
ment VIII, and from the latter, the new spe-
cies is distinguished by the hooked phallus
and lobate processes of the inferior append-
ages.
Distribution.—This species is known
499
only from the type locality in northern
Mexico.
Oxyethira desadorna Moulton and
Harris, new species
(Fig. 4)
Description.—Length 2.0 mm. Brown in
alcohol. 28 antennal segments. Segment VII
with short, acute, ventromesal process. Seg-
ment VIII subrectangular in lateral view,
posterior margin convex in middle; dorsal
margin with shallow, wide emargination;
ventral margin with deep, V-shaped exci-
sion. Segment IX complete dorsally and
concealed within segments VII and VIII; in
lateral view, anterior three-quarters wedge-
shaped, extending to middle of segment
VII. Tergum X fused with IX; nearly cir-
cular in dorsal view, posterior margin with
wide concavity. Subgenital plate thin and
elongate in lateral view, strongly curved
ventrad; in ventral view plate-like with
emarginate posterior margin, bearing pair
of setae posterolaterally. Inferior append-
ages fused; triangular in lateral view with
pair of internal processes, lower process
heavily sclerotized and upper process tu-
bular, bearing setae distally; posterior mar-
gin with broad, emargination in ventral
view, pair of setal bearing processes pro-
jecting from dorsal surface. Phallus tubular;
apex semimembranous and club-like, bear-
ing single, curved sclerotized process.
Immature stages and female-—Unknown.
Holotype male.—MEXICO; NUEVO
LEON, Municipio de Santiago, spring
along road above Cola de Caballo, 27 May
1991, S. Harris and A. Contreras (NMNH).
Paratypes—NUEVO LEON, Municipio
de Sanchez, Arroyo San Juan on road to
Laguna de Sanchez, 3.5 km W La Cienegra,
13 May 1989, S. Harris and A. Contreras,
56 36 (NMNH, INHS, SRM, SCH); TA-
MAULIPAS, Municipio de Gomez Farias,
Rio Frio at La Poza Azul, 6 km S Gomez
Farias, 7 August 1988, A. Contreras and A.
Moreno, 1 6 (NMNH).
Etymology.—Spanish; unadorned, refer-
ring to the simple appearance of the male
500 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
C
Fig. 4. Oxyethira desadorna, male genitalia. A, Left lateral view. B, Dorsal view. C, Ventral view. D, Phallus.
VOLUME 99, NUMBER 3
genitalia and the absence of distinctive ac-
cessories.
Diagnosis.—This species appears to be
most similar to O. unispina Flint and other
members of the Oxytrichia subgenus as de-
fined by Kelley (1984). As in O. unispina,
the phallus of O. desadoma has a narrow,
sinuate spine. However, the new species is
distinguished by its concave subgenital
plate and the absence of lateral extensions
from IX in lateral view.
Distribution Known only from the lo-
calities of the type material in northern
Mexico.
ACKNOWLEDGMENTS
We thank Joel Chirhart and Michael Pas-
anante for assisting with field collections of
caddisflies from East Texas and Kenneth
Stewart for providing travel support from
National Science Foundation grant DEB-
9347758. Oliver Flint and Gregg Easley
kindly provided us with additional speci-
mens of O. boquillas. Jon Gelhaus and
Ralph Garano provided collections and in-
formation based upon Big Bend National
Park permits BIBE-93-014 and BIBE-94-
019, respectively. Atilano Contreras and
Arnulfo Moreno were immense help to the
junior author during two collecting trips to
Mexico. Partial support for Contreras dur-
ing these trips was provided by the Univer-
sity of Alabama in 1989 and by the Uni-
versity of Minnesota Insect Collection in
501
1991. The Contreras-Ramos family is also
thanked for its hospitality during these col-
lecting trips. Allison Brigham, Gregg Eas-
ley, Jon Raese, and Dave Ruiter reviewed
earlier drafts of the manuscript. Comments
from Ralph Holzenthal and an anonymous
reviewer improved the quality of the final
draft.
LITERATURE CITED
Flint, O. S., Jr. 1972. Studies of Neotropical caddis-
flies, XIII: The genus Ochrotrichia from Mexico
and Central America (Trichoptera: Hydroptilidae).
Smithsonian Contributions to Zoology No. 118,
28 pp.
Harris, S. C. and J. L. Sykora. 1996. New species of
microcaddisflies from the eastern United States
(Insecta: Trichoptera: Hydroptilidae). Annals of
the Carnegie Museum 65: 17—25.
Holzenthal, R. W. and R. W. Kelley. 1983. New mi-
crocaddisflies from the southeastern United States
(Trichoptera: Hydroptilidae). Florida Entomolo-
gist 66: 464-472.
Kelley, R. W. 1984. Phylogeny, morphology and clas-
sification of the micro-caddisfly genus Oxyethira
Eaton (Trichoptera: Hydroptilidae). Transactions
of the American Entomological Society 110: 435—
463.
Marshall, J. E. 1979. A review of the genera of the
Hydroptilidae (Trichoptera). Bulletin of the British
Museum of Natural History (Entomology) 39:
135-239.
Morse, J. C. 1993. A checklist of the Trichoptera of
North America, including Greenland and Mexico.
Transactions of the American Entomological So-
ciety 119: 47-93.
Moulton, S. R., Il and K. W. Stewart. In press. A
preliminary checklist of the caddisflies (Trichop-
tera) of Texas. Proceedings of the Eighth Inter-
national Symposium on Trichoptera.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 502-504
A NEW GENUS AND A NEW SPECIES BELONGING TO THE SUBFAMILY
BLENNOCAMPINAE (HYMENOPTERA: TENTHREDINIDAE) FROM JAPAN
ICHIJI TOGASHI
1-chome, Honmachi, Tsurugi-machi, Ishikawa Prefecture 920-21, Japan.
Abstract.—Nipponostethus imperialis, n. gen., n. sp., from Japan are described and
illustrated. The new genus is closely related to Megatomostethus, and characters are given
to distinguish the two genera.
Key Words:
Recently, I studied eight specimens of
the subfamily Blennocampinae closely al-
lied to the genera Megatomostethus Tak-
euchi and Habachia Takeuchi. They differ
from these two genera by characters of the
malar space, the prepectus, the claw, and
the postorbital groove (Takeuchi 1952,
Okutani 1972). Therefore, I believe they
represent a new genus. In this paper, I de-
scribe and illustrate this new genus and spe-
CIES:
Nipponostethus Togashi, new genus
Body rather robust. Labrum rather small
(Fig. 3); front margin of clypeus slightly
emarginate (Fig. 3); malar space broad (Fig.
2); postorbital groove distinct (Fig. 2); post-
orbital carina present, developed below eye
(Fig. 2); antenna filiform, 3rd segment lon-
ger than 4th and segments 3—8 more than
2x longer than broad (Fig. 4); prepectus
present as raised shoulder, separated from
mesepisternum by furrow (Fig. 5); forewing
with 4 cubital cells, stub of analis straight
(Fig. 6); hindwing with a middle cell and
petiolate anal cell (Fig. 7); inner tibial spur
of fore leg cleft at apex (Fig. 8); tarsal claw
with inner tooth and basal lobe (Fig. 10).
Type species: Nipponostethus imperialis,
new species. This new genus is very closely
allied to the genus Megatomostethus Tak-
Nipponostethus, Blennocampinae, Tenthredinidae, sawfly, Japan
euchi, but it can be easily distinguished
from the latter by the presence of the broad
malar space (in Megatomostethus, the malar
space is linear or sometimes absent) and the
long slender antenna (in Megatomostethus,
the antenna is stout with segments 3-8 not
much longer than broad).
From Habachia Takeuchi, it is easily
separable by the small labrum (in Haba-
chia, the labrum is large and elongate); by
the presence of the prepectus (in Habachia,
the prepectus is absent); by the presence of
the large inner tooth and basal lobe of the
claws (in Habachia, the claw have a minute
inner tooth and no basal lobe); and by the
presence of the postorbital groove (in Ha-
bachia, the postorbital groove is absent).
Nipponostethus imperialis Togashi,
new species
(Figs. 1-12)
Female.—Length 7—7.5 mm. Head and
thorax black but apex of mandible reddish
brown; antenna entirely black; wings slight-
ly infuscate, stigma and veins black; legs
entirely black. Abdomen reddish brown,
with propodeum and sawsheath black.
Head: From above transverse (Fig. 1);
OOL:POL:OCL = 1.2:1.0:1.6; postocellar
area convex; interocellar and postocellar
furrows distinct but rather shallow; lateral
VOLUME 99, NUMBER 3
ae |
Figs. 1-11.
4
Nipponostethus imperialis. 1, Head, dorsal view. 2, Head, lateral view. 3, Clypeus and labrum,
front view. 4, Antenna, lateral view. 5, Mesopleuron, lateral view. 6, Forewing. 7, Hindwing. 8, Inner foretibial
spur, lateral view. 9, Hind tarsus, lateral view. 10, Tarsal claw. 11, Sawsheath, lateral view. Figs. 1, 2, 4, 5, 9,
scale = 1.0 mm; Figs. 3, 11, scale = 0.5 mm; Figs. 6, 7, scale = 5.0 mm; Figs. 8, 10, scale = 0.1 mm.
furrows distinct and deep (Fig. 1); frontal
area evenly concave, with a low surround-
ing wall; median fovea distinct, large, and
rather circular in outline; lateral foveae dis-
tinct and elongate; supraclypeal area rather
flattened; antenno-ocular distance slightly
longer than distance between antennal sock-
ets (ratio about 1.0:0.9). Clypeus convex;
labrum nearly flattened; malar space broad,
but nearly % as long as diameter of front
ocellus (ratio about 1.0:2.0).
Antenna slightly shorter than costa of
forewing (ratio about 1.0:1.1), relative
lengths of segments about 1.3:1.0:2.3:1.8:
eee 122112: pedicel longer than
width (ratio between length and width
about 1.0:0.7).
Thorax: Mesopraescutum prominently
raised; mesoscutellum nearly flattened; cen-
chri large, distance between them as long
as breadth of one. Wing venation as in Figs.
6 and 7. Hind basitarsus longer than follow-
ing 3 segments combined (ratio about 1.0:
0.8); inner fore tibial spur as in Fig. 8; claw
as in Fig. 10.
Abdomen: Normal; sawsheath as in Fig.
11; lancet with 16 serrulae (Fig. 12).
Punctation: Head except for clypeus and
labrum covered with fine setigerous punc-
tures; clypeus and labrum largely and rather
irregularly and closely punctured (Fig. 3);
pronotum, mesonotum and scutellum cov-
ered with fine setigerous punctures; poste-
rior margin of mesoscutellum closely and
striately sculptured; lateral sides of postter-
gite distinctly punctured; front and upper
portion of mesopleuron moderately and
rather closely punctured; mesosternum
nearly impunctate. Abdominal tergites sha-
greened.
Male.—Unknown.
Food plant—Unknown.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
=
Eve. 12:
Nipponostethus imperialis, lancet.
Habitat—Japan (Honshu and Shikoku).
Holotype.—Female, 18. IV. 1996, Impe-
rial Palace, Fukiage Gyoen, Tokyo Pref., A.
Shinohara leg. Deposited in the collection
of the National Science Museum (Nat.
Hist.), Tokyo.
Paratypes—3 @, 12. V. 1973, Mt. Ioo,
Ishikawa Pref., T. Mikage leg.; 1 2, 5. V.
1979, Mt. Kaji, Kochi Pref., T. Nishida leg.;
1 2, 24-26. VII. 1980, Minoto, Mts. Yat-
sugatake, Nagano Pref., A. Shinohara leg.;
1 2, 22. VII. 1989, Yarisawa (alt. 1600—
1900 m), Kamikochi, Nagano Pref., A. Shi-
nohara leg.; 1 2, 26. IV. 1992, Hikagezawa,
Mt. Takao, Tokyo Pref., A. Shinihara leg.
Two paratypes are deposited in the Smith-
sonian Institution, Washington, D.C., four
paratypes are deposited in the collection of
the National Science Museum (Natural His-
tory), Tokyo, and other one in my collec-
tion.
ACKNOWLEDGMENTS
I cordially thank Dr. David R. Smith,
Systematic Entomology Laboratory,
USDA, Washington, D.C., for his kind ad-
vice and review of the manuscript. I am in-
debted to Dr. A. Shinohara, National Sci-
ence Museum (Nat. Hist.), Tokyo, Mr. T.
Mikage, Ageo City, Saitama Pref., and Mr.
T. Nishioka, Kochi City, Kochi Pref., for
giving me the valuable specimens.
LITERATURE CITED
Okutani, T. 1972. A new genus and a key to Japanese
Genera of the subfamily Blennocampinae (Hym.
Tenth.). Entomological Review, Japan 24: 57-61.
Takeuchi, K. 1952. A generic classification of the Jap-
anese Tenthredinidae (Hymenoptera: Symphyta).
Kyoto. 90 pp.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 505-512
A FIRST RECORD OF APTILOTUS MIK (DIPTERA: SPHAEROCERIDAE)
FROM THE NEOTROPICAL REGION, WITH THE DESCRIPTION OF
THREE NEW WINGLESS SPECIES OF THE APTILOTUS PARADOXUS
GROUP FROM HIGH ELEVATIONS IN COSTA RICA
S. A. MARSHALL
Department of Environmental Biology, University of Guelph, Guelph, Ontario, Canada
N1G 2WI1.
Abstract.—The genus Aptilotus Mik is recorded from the Neotropical Region for the
first time. Aptilotus nigritibia, new species, A. nigrimera, new species, and A. zumbadoi,
new species, all completely wingless, are described from ca. 3,000 m in Costa Rica. A
key to the Costa Rican Aptilotus is provided.
Key Words:
Costa Rica
The genus Aptilotus, named for the sin-
gle wingless European species, A. para-
doxus Mik, is a primarily Holarctic group
in which wing reduction has occurred in-
dependently within at least five lineages
(Marshall 1983, Marshall and Smith
1990). Twelve described species of Aptil-
otus are Nearctic, one species is described
from Europe, one is described from Ethi-
opia, four are from Nepal, two are from
southeast Asia, and a group of seven
closely related species are restricted to the
Canary Islands. There were no known
Neotropical Aptilotus prior to the discov-
ery of the three new wingless species de-
scribed herein, all of which are endemic
to high elevation sites in Costa Rica.
Aptilotus species can be apterous, bra-
chypterous, or polymorphic for wing de-
velopment, with aptery having evolved
independently in each of the species
groups recognized by Marshall (1983).
The genus can be diagnosed by the fol-
lowing characters: male with a translu-
cent, usually quadrate posteromedial lobe
Diptera, Sphaeroceridae, Limosininae, wing loss, zoogeography, taxonomy,
on sternite 5; body usually black, shining
and punctate; female tibia with only an
apical bristle; dorsocentral bristles in 2
pairs but anterior pair often small; sursty-
lus of characteristic shape, divided into an
outer setose part and an inner part with 1
or (usually) 2 stout bristles.
All known Central American Aptilotus
can be distinguished from other wingless
Central American Sphaeroceridae not
only by the defining characters of Apftil-
otus as outlined above, but also by the
plesiomorphic possession of two orbital
bristles. Other wingless Central American
sphaerocerids, most of which belong to
the genus Pterogramma, have only a sin-
gle orbital bristle.
RELATIONSHIPS
The species described here belong to
the paradoxus group of Marshall (1983)
with which they share the following syn-
apomorphies: distiphallus elongate, com-
prised of distinctive dorsal and ventral
loops and a broad, bilobed membranous
506
and microspinulose apex; outer posterior
part of surstylus long-setose; anterior sur-
stylar spur characteristically flattened, of-
ten scale-like. Delimited on the basis of
these characters, the paradoxus group in-
cludes a clade of 6 brachypterous and one
macropterous species restricted to the Ca-
nary Islands (A. beckeri and related spe-
cies), an apterous European species (A.
paradoxus Mik), three macropterous spe-
cies from Nepal (A. glabrifons Marshall,
A. spinistylus Marshall, A. rufiscapus
Marshall), two macropterous eastern
North American species (A. pogophallus
Marshall and A. nigriphallus Marshall), a
macropterous species in Japan (A. longi-
nervis Hayashi), a dimorphic western
North American species (A. /uctuosus
(Spuler)) and two brachypterous western
North American species (A. nigriscapus
Marshall, A. luteoscapus Marshall). The
new Costa Rican species all seem to be
closely related to a clade including Eu-
ropean and western North American spe-
cies which have relatively short female
cerci and quadrate, bilobed posteromedial
areas on male sternite 5.
Although all Aptilotus exhibit an unusual
degree of morphological uniformity, the
Costa Rican species seem to represent a dis-
tinct clade, weakly supported on the basis
of the shape of sternite 5 of the male, the
sclerotized part of which is very short me-
dially, and sternite 8 of the female, which
is reduced and modified. Within the Costa
Rican Aptilotus clade, nigrimera plus zum-
badoi form a monophyletic group, strongly
supported by the possession of a third sur-
stylar bristle and the loss of bristles from
tergite 9 of the female.
DISCUSSION
All of the species described here were
collected on Cerro de la Muerte, at a cold
paramo-like site at 3300 m, or in high el-
evation forests (2600 m and above) within
a few kilometres of Cerro de la Muerte.
Although it seems likely that related spe-
cies occur, or previously occurred, at oth-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
er relatively high elevation sites in the in-
tervening area, current data show a dis-
junction between the cluster of endemic
Aptilotus in Costa Rica and their nearest
relatives in western North America. Other
high elevation collections in Costa Rica
have yielded wingless Sphaeroceridae,
but only in primarily Neotropical clades
related to Pterogramma Spuler. Similarly,
sphaerocerid samples available from leaf
litter collections from other high-eleva-
tion Central American sites have included
a diversity of undescribed Pterogramma
but no Aptilotus. Habitat at these sites
ranged from mixed oak-pine forests at
2800 m (Oaxaca, Mexico) to high eleva-
tion cloud forests at elevations from
2300—2800 m (Chiapas, Mexico; Quezal-
tenango, Guaemala; El] Salvador; Cerro
Chiriqui, Panama). Also, no South Amer-
ican Aptilotus are known, despite collec-
tion efforts in apparently suitable habitat
in the paramos of Ecuador and Venezuela.
Despite the lack of Mexican and Cen-
tral American records outside Costa Rica,
further sampling at higher elevations than
those sites from which samples are cur-
rently available are likely to reveal addi-
tional Aptilotus species. It is hypothesized
that Aptilotus is a primarily northern
group, present in the Neotropical region
as far south as lower Central America (but
not into South America), where it exhibits
a highly disjunct distribution restricted to
cool temperate conditions at elevations
above 2500 m.
KEY TO THE COSTA RICAN SPECIES OF
APTILOTUS
1. Legs mostly black, trochanters and mid and
hind tarsi brown; antenna black. Female ster-
nite 8 very small, transverse (Fig. 14). Female
tergite 9 with two bristles (Fig. 16). Surstylus
with 2 short, thick ventral bristles (Fig. 11)
Pe One ORIG Ons eee Aptilotus nigritibia, n. sp.
— Legs luteous, antenna variable. Female sternite
8 with an elongate anterior portion (Figs. 6,
22). Female tergite 9 bare (Fig. 8). Surstylus
with 3 short, thick ventral bristles (Figs. 4, 20).. 2
VOLUME 99, NUMBER 3
2. Antenna black, legs mostly luteous. Part of
frons and body bare and shining. Middle ven-
tral surstylar bristle inserted close to postero-
ventral surstylar bristle (Figs. 1, 4) .......
SE Ts ae er 0, Aptilotus nigrimera, 0. sp.
— Antenna and legs mostly luteous. Frons and
body entirely pruinose, without shining areas.
Middle ventral surstylar bristle mid way be-
tween anteroventral and posteroventral surstylar
bristle (Figs. 17, 20) .. Aptilotus zumbadoi, n. sp.
DESCRIPTIONS OF COSTA RICAN APTILOTUS
Aptilotus nigrimera Marshall, NEw
SPECIES
(Figs. 1-8)
Description.—Body length 2.5—3.0mm.
Body shining black; fore tarsus pale brown,
legs otherwise yellow; antenna black to
dark brown. Middle part of frons mostly
bare and shining, ocellar triangle setulose
but flanked by bare areas, interfrontal area
with only a few setulae medially. Interfron-
tal bristles in 2—3 thin, subequal pairs. Eye
height 4.0 genal height. Dorsocentral bris-
tles in two postsutural pairs, anterior pair
only slightly longer than acrostichal setulae,
prescutellar pair long; only 2—3 rows of ac-
rostichal setulae between dorsocentral ar-
eas. Scutellum almost transverse, twice as
wide as long. Mid tibia with a long anter-
odorsal bristle proximally, a long distal dor-
sal bristle, and short anterodorsal and pos-
terodorsal distal bristles. Ventral surface of
tibia with only an apical bristle in both sex-
es. Wing and halter completely absent.
Male abdomen.—Tergite 1+2 longer
than other tergites, uniformly dark. Sternite
5 with anterior and posterior margins
strongly curved, anterior margin with with
shallow medial notch; posteromedial pale
part of sternite extending back to anterior
margin of sternite so that pigmented part of
sternite is very short medially, pale poster-
omedial section with brown pigmentation
medially, rounded and narrowly notched
posteromedially; pale area flanked by clus-
ters of bristles. Surstylus broad, with thin
bristles on a posterolateral bulge or ridge
and 3 thick ventral bristles; a flattened,
broadly bifid anteroventral bristle, a short,
507
thick posteroventral bristle and a long, thick
ventral bristle inserted much closer to pos-
teroventral than anteroventral bristle. Gon-
ostylus almost parallel sided, thick, distal
part tapered. Basiphallus short, quadrate.
Distiphallus with dorsal (functionally ven-
tral) sclerite with a broad basal part, a nar-
row intermediate section and a distally
forked part; ventral sclerite with a broad
base and broad distal loop; membranous
part of distiphallus with long distal dorsal
lobes covered with flattened setulae.
Female abdomen.—Tergite 8 black, long,
with a shallow anterior notch. Tergite 9
pale, with a short, transverse, anterior part
and a large posterior part narrowly fused
with cerci, surface bare (the usual 2 bristles
absent). Cerci pale, shorter than tergite 9.
Sternite 8 long and narrow, posterior part
slightly enlarged with 4 small bristles. Ster-
nite 9 broad, setulose except lateral ex-
tremes. Two internal vaginal sclerites pres-
ent in addition to large, thin-rimmed rings
joined by a discontinuous transverse piece
(spectacles-shaped sclerite of Rohdcek,
1983). Spermathecae spherical, sclerotised
parts of ducts slightly longer than sperma-
thecal body.
Types.—Holotype (¢,INBio) and 5 para-
types (3¢,22,GUE): COSTA RICA. Car-
tago Province, km. 89 Highway 2, Cerro de
la Muerte, 10.11.1995, aspirated among
bamboo litter, S.A. Marshall. Paratype
(36,GUE) same as above, but sifted by R.S.
Anderson.
Etymology.—The specific epithet refers
to the dark antenna, a diagnostic character
separating A. nigrimera from the closely re-
lated A. zumbadoi.
Aptilotus nigritibia Marshall, NEw
SPECIES
(Figs. 9-16)
Description.—Body length 2.0—2.5mm.
Body almost entirely heavily pruinose,
black; fore tibia and tarsi black, other tarsi
brown, legs otherwise yellow; antenna yel-
low to pale brown. Head entirely pruinose,
without shining areas. Interfrontal bristles
508 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-8. Aptilotus nigrimera. 1, Male terminalia, left lateral. 2, Aedeagus and associated structures,
left lateral. 3, Distiphallus, ventral view. 4, Surstylus, ventral view. 5, Sternite 5 of male. 6, Female ter-
minalia, ventral. 7, Spermathecae and associated sclerites. 8, Female terminalia, dorsal. Abbreviations: gs
= gonostylus (paramere of earlier papers); T9 = tergite 9 (epiproct of earlier papers).
in 2—3 thin, subequal pairs. Eye height 4.5 x
genal height. Dorsocentral bristles in two
postsutural pairs, anterior pair twice as long
as acrostichal setulae, prescutellar pair
much longer; 4 rows of acrostichal setulae
between dorsocentral areas. Scutellum al-
most transverse, twice as wide as long. Mid
tibia with a long anterodorsal bristle prox-
VOLUME 99, NUMBER 3
509
Figs. 9-16. Aptilotus nigritibia. 9, Male terminalia, left lateral. 10, Aedeagus and associated structures, left
lateral. 11, Distiphallus, ventral view. 12, Surstylus, ventral view. 13, Sternite 5 of male. 14, Female terminalia,
ventral. 15, Spermathecae and associated sclerites. 16, Female terminalia, dorsal.
imally, a long distal dorsal bristle, a short
anterodorsal distal bristle and a smaller pos-
terodorsal distal bristle. Ventral surface of
tibia with only an apical bristle in both sex-
es. Wing and halter completely absent.
Male abdomen.—Tergite 1+2 longer
than other tergites, uniformly dark. Sternite
5 with anterior and posterior margins
strongly curved, anterior margin with shal-
low anteromedial notch; posteromedial pale
part of sternite extending back to anterior
margin of sternite so that pigmented part of
sternite is very short medially; pale poster-
omedial section with brown pigmentation
medially, rounded and narrowly notched
posteromedially; pale area flanked by clus-
510
ters of bristles. Surstylus broad, with thin
bristles on a posterolateral bulge or ridge
and 2 thick ventral bristles; a flattened an-
teroventral bristle and a short, thick pos-
teroventral bristle; a very small ventral bris-
tle inserted close to posteroventral bristle.
Gonostylus of medium width and weakly
S-shaped, distal part expanded. Basiphallus
short, quadrate. Distiphallus with dorsal
(functionally ventral) sclerite with a broad
basal part, a narrow intermediate section
and a distal loop which is not closed dis-
tally; ventral sclerite with a broad base and
broad distal loop; membranous part of dis-
tiphallus with long distal dorsal lobes cov-
ered with conspicuous flattened setulae.
Female abdomen.—Tergite 8 black, long,
with a shallow anterior notch. Tergite 9
pale, with a short, transverse, anterior part
and a large posterior part narrowly fused
with cerci and with 2 dorsal bristles. Cerci
pale, shorter than tergite 9. Sternite 8 small,
transverse, with 4 small bristles. Sternite 9
broad, setulose except lateral extremes. In-
distinct vaginal sclerites present in addition
to large, thin-rimmed rings joined by a dis-
continuous transverse piece (spectacles-
shaped sclerite of Rohacek 1983). Sper-
mathecae spherical, sclerotised parts of
ducts slightly longer than spermathecal
body.
Types.—Holotype (6 ,INBio) and 2 para-
types (1d,12,GUE): COSTA RICA: Car-
tago Province, km. 89, Highway 2, Cerro
de la Muerte, 10.11.1995, aspirated among
bamboo litter, S.A. Marshall.
Etymology.—The specific epithet refers
to the black tibia which differentiates A. ni-
gritibia from Costa Rican congeners.
Aptilotus zumbadoi Marshall, NEw
SPECIES
(Figs. 17—24)
Description.—Body length 2.0—2.5mm.
Body almost entirely heavily pruinose,
black; fore tibia and tarsi black, other tarsi
brown, legs otherwise yellow; antenna yel-
low to pale brown. Head entirely pruinose,
without shining areas. Interfrontal bristles
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
in 2—3 thin, subequal pairs. Eye height 4.5 x
genal height. Dorsocentral bristles in two
postsutural pairs, anterior pair twice as long
as acrostichal setulae, prescutellar pair
much longer; 4 rows of acrostichal setulae
between dorsocentral areas. Scutellum al-
most transverse, twice as wide as long. Mid
tibia with a long anterodorsal bristle prox-
imally, a long distal dorsal bristle, a short
anterodorsal distal bristle and a smaller pos-
terodorsal distal bristle. Ventral surface of
tibia with only an apical bristle in both sex-
es. Wing and halter completely absent.
Male abdomen.—Tergite 1+2 longer
than other tergites, uniformly dark. Sternite
5 of uniform length, neither anterior nor
posterior marging strongly curved, anterior
margin with broad medial notch; postero-
medial pale part of sternite extending back
almost to anterior margin of sternite, with
elongate brown pigmentation medially,
quadrate and narrowly notched posterome-
dially; pale area flanked by clusters of bris-
tles. Surstylus broad, with thin bristles on a
posterolateral bulge or ridge, and 3 thick
ventral bristles; a flattened anteroventral
bristle, a short, thick posteroventral bristle
and a ventral bristle midway between the
other two. Gonostylus thin, of uniform
width and weakly S-shaped. Basiphallus
short, quadrate. Distiphallus with dorsal
(functionally ventral) sclerite with a broad
basal part, a narrow intermediate section
and a distal loop; ventral sclerite with a
broad base and broad distal loop.
Female abdomen.—Tergite 8 long, bare
and shining medially and anteriorly, with a
shallow anterior notch. Tergite 9 pale, with
a short, transverse, anterior part and a large
posterior part narrowly fused with cerci,
surface bare (the usual 2 bristles absent).
Cerci slightly longer than tergite 9. Sternite
8 long and narrow, posterior part separate
from long and narrow part, with 4 small
bristles. Sternite 9 broad, setulose except
lateral extremes. Indistinct internal vaginal
sclerotisation present in addition to large,
thin-rimmed rings joined by a broad sclerite
(spectacles-shaped sclerite of Rohdcek
VOLUME 99, NUMBER 3
511
Figs.17—24. Aptilotus zumbadoi. 17, Male terminalia, left lateral. 18, Aedeagus and associated structures,
left lateral. 19, Distiphallus, ventral view. 20, Surstylus, ventral view. 21, Sternite 5 of male. 22, Female ter-
minalia, ventral. 23, Spermathecae and associated sclerites. 24, Female terminalia, dorsal.
1983). Spermathecae spherical, sclerotised
parts of ducts slightly longer than sperma-
thecal body.
Types.—Holotype (¢, INBio) and 3
paratypes (26,12,GUE): COSTA RICA:
Cartago Province, km. 89, Highway 2, Cer-
ro de la Muerte, 10.11.1995, aspirated
among bamboo litter, S.A. Marshall.
Paratypes.—COSTA RICA. Cartago
Province, 2kmS Villa Mills, 3000m,
10.11.1996, ridge top mature oak forest litter,
R.S. Anderson (1¢,22,GUE); San Jose
Province, km. 68 Highway 2, Tres de Junio
Bog, 2600m, 10.11.1996, litter from forest
adjacent to Sphagnum bog, R.S. Anderson
(236,GUE); Cerro de la Muerte, 7-—
13.iv.1985, pan traps, oak cloud forest, L.
Masner and H. Goulet (1¢,GUE); km. 95,
Highway 2, 3200m, 13.iv.1985, oak cloud
forest, L. Masner (1¢, GUE).
S12
Etymology.—The specific epithet is a pa-
tronym in recognition of INBio’s Diptera
specialist, Manuel Zumbado. Without Man-
uel’s help we could not have made collec-
tions at the type localities for this and other
species described here.
ACKNOWLEDGMENTS
Dr. Robert Anderson, Canadian Museum
of Nature, provided specimens, advice con-
cerning high altitude insects of Central
America, and company in the field. Manuel
Zumbado, Instituto Nacional de Biodiver-
sidad (INBio), Costa Rica, provided invalu-
able advice and field support. Rebecca
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Langstaff did the illustrations. Paratypes are
retained in the University of Guelph Insect
Collection (GUE).
LITERATURE CITED
Marshall, S. A. 1983. A revision of the genus Apztil-
otus Mik in North America (Diptera; Sphaerocer-
idae). Canadian Journal of Zoology 61: 1910-—
1924.
Marshall, S. A. and I. P. Smith 1990. A review of the
North American species of Aptilotus, with de-
scriptions of new species from North America and
Nepal (Diptera; Sphaeroceridae). Canadian Jour-
nal of Zoology 68: 2338-2351.
Rohaéek, J. 1983. Monograph and reclassification of
the previous genus Limosina Macquart (Diptera,
Sphaeroceridae) of Europe. Beitrage zur Ento-
mologie, Berlin 33: 3-195.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 513-522
COMPARISON OF FIXATION AND DRYING PROCEDURES FOR SCANNING
ELECTRON MICROSCOPY AMONG INSECT BODY TYPES
MICHEL SWEARINGEN, DAVID HEADRICK, AND TOM BELLOWS
Department of Entomology, University of California Riverside, CA 92521, U.S.A.
Abstract.—Four different procedures used to prepare insect specimens for scanning
electron microscopy were compared using three insect body types that represent unique
obstacles to obtaining good quality micrographs, 1.e., clean and life-like in appearance.
The micrographs produced using each procedure were rated for cleanliness, structural
integrity, and imaging quality. Each procedure involved: fixation, post-fixation, and dry-
ing. Procedure 1 involved fixation by freezing @ —10°C and drying by critical-point
drying. Procedure 2 involved fixation by submersion in 70% ethanol and drying by crit-
ical-point drying. Procedure 3 involved fixation by submersion in 70% ethanol and drying
with hexamethyldisilizane. Procedure 4 involved fixation by submersion in a hot water
bath at 80°C and drying with hexamethyldisilizane. Procedure 2 gave the best overall
results, but procedures 3 and 4 were best on insects with small lightly sclerotized bodies
and unsclerotized larvae, respectively. Difficulties associated with histological study of
insects are discussed.
Key Words:
fixation, procedures
Scanning electron microscopy (SEM) is
used for a variety of purposes in Entomol-
ogy. The most common applications in-
volve taxonomic and morphological stud-
ies. While SEM is often an indispensable
tool in these types of studies, the quality of
the images can vary considerably; much of
this variation is related to problems specific
to insects. Quality images useful for scien-
tific study require that the specimens be
clean and life-like in appearance (i.e. little
or no shrinkage).
The procedures for preparation of insect
specimens for SEM are fairly standard, al-
though variations during fixation and de-
hydration are common (see below). Papers
that include SEM micrographs may not de-
scribe their procedures or simply acknowl-
edge an individual or department for their
help in the production of SEM micro-
Scanning electron microscopy, freezing fixation, ethanol fixation, hot water
graphs, thus, information on new or differ-
ent successful techniques are difficult to ob-
tain. Insects present a wide range of histo-
logical variables, and no one treatment is
universally applicable to every taxa or
stage. The objective of this study was to
compare different procedures used on dif-
ferent insect body types to determine
which, if any, provided consistent and pre-
dictable results, or which procedure was
best suited with selected taxa to obtain
clean, life-like specimens.
For the purposes of this paper, we assume
the reader has basic knowledge of SEM
preparation methods; however, the follow-
ing terms are defined:
Accelerating voltage.—Accelerating
voltages are typically set at ca. 10—20kV
for biological tissues. The resultant resolu-
tion of surface detail usually does not ex-
ceed 10,000 diameters of magnification.
514
Fixation.—The cessation of life in such
a manner as to retain the structural and bio-
chemical integrity of the specimen and its
tissues. See Sabatini et al. (1963), Human-
son (1967), Barbosa (1974), and Dawes
(1988) for basic principles of fixation. This
is the most important step in histological
preparation.
Postfixation.—There are many postfixa-
tion processes for biological tissues depend-
ing on the examination method. Metallic
impregnation with heavy metal salts is
commonly used for SEM (Sabatini, et al.
1963, Dawes 1988). The most common me-
tallic salt is osmium tetroxide (OsO,) which
acts to bind lipids, thus increasing the elec-
tron density of the tissues. Electron density
in biological tissues allows for higher ac-
celerating voltages without charging, with
the subsequent gain in resolution. Enhanced
contrast of the specimen also can result
from metal impregnation leading to a better
quality image for study (Dawes 1988).
OsO, is labeled as a poison. A Material
Safety Data Sheet is provided with the pur-
chase of OsO,; precautions on proper han-
dling and disposal should be followed.
Drying (Dehydration).—Tissues must be
completely dry, including absence of met-
abolic water, before sputter-coating and ex-
amination with SEM. There are several dry-
ing procedures. The most common is the
use of a critical-point dryer with liquid CO,
as the transition fluid (Dawes 1988, Gordh
and Hall 1979). Hexamethyldisilizane
(HMDS) is another chemical which dries
tissues (Adams et al. 1987, Nation 1983)
and is employed in some of the procedures
listed below. HMDS is labeled as corrosive
and highly toxic. Again, a Material Safety
Data sheet is provided with purchase and
procedures for proper handling should be
followed.
Body types.—Three distinct body types,
difficult to successfully prepare for SEM,
were selected for analysis A) small bodies
(<2 mm long), B) bodies that produce wax
and C) soft-bodied forms, which include
both immature stages or insect imagos and
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
immatures with little sclerotization of the
integument. These types are referred to as
‘small’, ‘waxy’ or ‘soft’ body types. We
used Encarsia sp. (Hymenoptera: Aphelin-
idae) a parasitoid that is ca. 1.0 mm long,
to represent the small body type. For waxy
body types, we used the nymphs of two
species of whitefly, Bemisia argentifolii
Bellows and Perring and Dialeurodes citri
(Ashmead) (Homoptera: Aleyrodidae). The
nymphs of these two whitefly species also
are soft-bodied. We used late-instar larvae
of Musca domestica L. (Diptera: Muscidae)
to represent the soft body type.
SPECIMEN PREPARATION
The problems associated with insects and
their preparation for SEM involve primarily
the cuticle, and secondarily, the presence of
waxes and lipids. All insects possess a mul-
ti-layered cuticle which presents a unique
obstacle to histology and, depending on the
taxa involved, may be heavily sclerotized
or rather thin and flexible (Humason 1967,
Barbosa 1974).
The cuticle provides a barrier to the ma-
terials used in each step in specimen prep-
aration: fixation, postfixation and drying
(Barbosa 1974, Dawes 1988). During fixa-
tion, the cuticle can prevent adequate pen-
etration of the fixative into the body, and
thorough fixation of tissues. Poor fixation
may also cause problems later during post-
fixation as tissues that have not been fixed
will not accept the post-fixative, and may
be more susceptable to electron absorption
which results in charging (Dawes 1988).
Drying of the tissues may also be hampered
by partial fixation, allowing water and al-
cohol to remain in the specimen. Such tis-
sues may burst when placed in a vacuum
during sputter-coating and SEM examina-
tion. Metamorphosis also poses special
problems during processing of tissues for
SEM, for if the specimen being processed
is in a period of structural rearrangement,
the cuticle may be prone to shrinkage and
severe charging problems.
Other problems associated with SEM ex-
VOLUME 99, NUMBER 3
amination of insects involve the presence of
waxes and lipids. All biological tissues con-
tain lipids, which tend to absorb electrons,
and thus lead to charging. Many insects
also produce waxes which also do not fix
well, absorb electrons, and may obscure
certain structures. The post-fixative, osmi-
um tetroxide, is typically used to bind lipids
to make them more electron dense, thus
eliminating electron absorption and conse-
quent charging (Dawes 1988).
SEM facilitates examination of the sur-
face features of very tiny structures. Many
insects are quite small or bear minute struc-
tures on their bodies. Although easily pen-
etrated by preparation materials, these
small-bodied insects pose difficulties during
examination as they tend to become elec-
trically charged.
With these problems in mind, we applied
four procedures to specimens of taxa which
represent the three body types described
above. Comparisons of the procedures are
presented with a rating scale for each.
MATERIALS AND METHODS
Each procedure involved three basic
steps: 1) fixation, 2) post-fixation, and 3)
drying. Three methods of fixation were em-
ployed in the current study: 1) freezing @
—10°C, 2) submersion in ethanol (50—-
70%), and 3) a hot water bath at 80°C. A
2% aqueous solution of osmium tetroxide
was always used as the postfixative. Metal-
lic impregnation as a postfixative may not
be commonly used for some insect taxa, but
has been shown to be beneficial and thus is
included in each procedure (Dawes 1988,
Headrick, unpublished data). Dehydration
involved placing the specimens in a series
of ethanol baths at dilutions of 15, 30, 50,
70, 80, 90, 95 and finally 100%, followed
either by critical-point drying using liquid
CO, as a transition fluid, or HMDS. Fol-
lowing the 100% ethanol bath in two pro-
cedures the specimens were placed in
HMDS under a fumehood. After 20 min.
the remaining HMDS was siphoned away
with a pipette, thus removing any accum-
Syl)
lated particles or residues. A second bath of
clean HMDS was added and the specimens
were dried to completion by evaporation of
the HMDS in a fumehood (ca. 30 min).
Procedure 1.—This method used freez-
ing @ —10°C for fixation. The specimens
were postfixed by immersion in the solution
of osmium tetroxide for 24 h. They were
next rinsed twice with deionized water, then
dehydrated through a series of increasing
ethanol dilutions. Specimens remained in
each dilution for 20 min and ended in 100%
ethanol. Drying used critical-point drying.
This procedure was applied to 10 speci-
mens each of Dialeurodes citri, Bemisia ar-
gentifolii, and Encarsia sp.
Procedure 2.—Specimens were fixed in
50% ethanol for 24 h, then rehydrated to
distilled water through a decreasing series
of ethanol dilutions for post-fixation in os-
mium tetroxide. Postfixation, dehydration,
and drying were identical to procedure 1.
Procedure 2 was applied to 10 specimens
each of D. citri, B. argentifolii, and E. sp.
Procedure 3.—Fixation was in 70% eth-
anol, and postfixation in 2% osmium te-
troxide for 24 h. The specimens were then
dehydrated through a series of increasing
ethanol dilutions up to 100% for 15 min in
each dilution. The specimens were dried us-
ing two, 30-min baths of HMDS. This pro-
cedure was applied to 10 specimens of E.
sp.
Procedure 4.—Specimens were fixed in a
bath of 80°C tap water and subjected to ul-
trasonic cleaning for 10 min to remove res-
idue from the body (Belcari 1987). After 5
min of fixation, the specimens were cut in
half to facilitate the fixation process. The
specimens were then post-fixed in osmium
tetroxide for 24 h, and rinsed twice with
deionized water. The specimens next were
dehydrated in an increasing series of etha-
nol dilutions, up to 100%; each ethanol bath
lasted 20 min. Two 30 min baths in HMDS
finally were used to dry the specimens. This
procedure was applied to 12 specimens of
M. domestica.
All dried specimens were mounted on
516 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
aluminum stubs with double-sided cello-
phane tape and sputter-coated with a gold-
palladium alloy using a Hummer V. Speci-
mens were examined on a JEOL JSM C35
scanning electron microscope in the De-
partment of Nematology at the University
of California, Riverside. Scanning electron
micrographs were prepared at 15 kV accel-
erating voltage on Polaroid 55P/N film.
To quantify the results of our compara-
tive survey, we used the following rating
system. We judged SEM micrograph qual-
ity on three variables: cleanliness, structural
integrity, and imaging. Each of these cate-
gories had two subdivisions. We rated
cleanliness at two levels, “‘particle’’ and
“obscuring.” Particle is macroscopic ac-
cumulation of atmospheric or environmen-
tal debris by the specimen. Particle accu-
mulation can take place during any phase
of the processes used for specimen prepa-
ration. Obscuring is a microscopic phenom-
enon in which materials such as waxes, res-
ins or other natural products are chemically
affected by the SEM preparation processes
and adhere to the surface of the specimen,
thus obscuring minute surface details. Sim-
ilar obscuring problems arise from metallic
filming or sputter-coating processes (Dawes
1988). The rating system ranged from 0-3
in both categories. For particle cleanliness,
a 0 indicated that the specimen was covered
with particles, and thus rendered useless for
study (Fig. 1A); while 3 meant the speci-
men was free of any obscuring particles.
With respect to obscuring residues, a 0
meant minute structures were obscured
from view, and a 3 indicated that all struc-
tures could be seen clearly.
Structural integrity includes both macro-
scopic and microscopic levels. Macroscopic
structural integrity encompasses the body as
a whole. Microscopic structural integrity re-
lates to minute structures on the body, such
as setae or sensoria. We rated both of these
categories as good, fair or poor. Good struc-
tural integrity evidenced little to no struc-
tural collapse. Fair structural integrity may
have some collapse, but not loss of useable
micrographs for study. Poor structural in-
tegrity resulted from collapse of the speci-
men, or loss of small structural details, pre-
venting observation of important features
ist Bye):
Imaging includes resolution and charg-
ing. Resolution at magnifications greater
than 2000 was rated as 0-3. A 0 indicated
that the image could not be adequately re-
solved and a 3 indicated that resolution
above 10,000 was obtainable. Charging in
specimens is either present (+) or absent
(—) (Fig. 1D).
Our rating system is summarized as fol-
lows and was used to build Tables 1-3:
A. Specimen cleanliness:
1. Particle—O-3
2. Obscuring—0-3
B. Structural Integrity:
1. Macroscopic—Good, fair, or poor
2. Microscopic—Good, fair, or poor
C. Imaging:
1. Resolution at higher magnifications
(>2000): 0-3
2. Charging: present (+) or absent (—).
RESULTS
Procedure 1.—Freezing fixation
Dialeurodes citri.—With this procedure
specimens were generally clean and no
structures were obscured from view (Fig.
2A). At higher magnification, resolution
was rated as 1 (Fig. 2B). Collapse of the
body and its smaller structures was not ob-
served and charging did not occur with
these specimens.
Bemisia argentifolii.mCleanliness with
respect to particles was good (Fig. 2C), but
most minute structures were obscured by
waxes. Because the specimens were not
clean, the tracheal furrows were left full of
wax (Fig. 2D); this was more life-like, but
obscured potentially important taxonomic
characters. Severe charging was present in
some specimens. Resolution at higher mag-
nification was rated as 1.
Encarsia sp.—The specimens bore ob-
scuring residues (Fig. 2E). Macroscopic and
a
VOLUME 99, NUMBER 3
LokW
Fig. 1.
gei4
Examples of poor SEM micrographs. A, Surface details obscured by particulate matter, this should
look like Figure 5C. B, Shinkage of soft-bodied dipteran larva. The shrinkage on this specimen was so severe
that even the most minute of structures was collapsed. C, Shinkage of a hymenopteran parasitoid antenna. D,
An example of what charging does to a micrograph.
Table 1. Rating results for Procedure 1, freezing
fixation.
Taxa
Rating D. citri B. argentifolii ESSSp:
Cleanliness
Particle 2 2 2
Obscuring 2 1
Structural integrity
Macro good good poor
Micro good fair poor
Imaging
Resolution l O
Charging (—) (+) Ga)
Table 2. Rating results for Procedure 2, ethanol fix-
ation.
Taxa
Rating D. citri B. argentifolii E. sp:
Cleanliness
Particle 2 2 2
Obscuring 2) 2 2
Structural integrity
Macro good good good
Micro good good good
Imaging
Resolution 2 2
Charging (j) G) (|)
518
Table 3. Rating results for Procedures 3, ethanol
fixation and HMDS, and Procedure 4, water bath fix-
ation and HMDS.
Procedure 3 Procedure 4
Rating E. sp. M. domestica
Cleanliness
Particle 2) 3
Obscuring 3) 3
Structural integrity
Macro fair good
Micro good good
Imaging
Resolution 3 3
Charging (Ge) ()
microscopic structures showed considerable
collapse (Fig. 2F). No charging occurred,
but resolution at higher magnifications was
rated as 0.
Procedure 2.—Ethanol fixation
Dialeurodes citri.—The specimens were
free of particles and obscuring residues
(Fig. 3A). Resolution at higher magnifica-
tion was rated as 2 (Fig. 3B). Collapse and
charging were not problems.
Bemisia argentifolii.imThe specimens
(Fig. 3C), and specifically, the tracheal fur-
rows were free of waxes (Fig. 3D). Reso-
lution was rated as 2 at higher magnifica-
tion. There was no structural collapse or
charging.
Encarsia sp.—The specimens were clean
with little collapse (Fig. 3E). At higher
magnifications, the resolution obtained was
rated as 1 (Fig. 3F).
Procedure 3.—Ethanol fixation, HMDS
Encarsia sp.—The specimens were most-
ly free of particles or obscuring residues
(Fig. 4A). Microscopic collapse of struc-
tures was not a problem with this procedure
(Fig. 4B, C); although there was some mac-
roscopic collapse in the area of the propo-
deum (Fig. 4A). Resolution was extremely
good up to 18,000 (Fig. 4C), and there
was no charging.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Procedure 4.—Hot water fixation/
sonication, HMDS
Musca domestica.—The specimens were
very clean with minimal obscuring residues
(Fig. 5A, B, C). No collapse was evidenced
and the smallest structures remained life-
like in appearance (Fig. 5C). Resolution
was excellent and there was no charging.
DISCUSSION
Casual examination of entomological
journals reveals a considerable range in the
quality of SEM micrographs. Technological
advances in SEM’s have made them easier
to use and have provided the user with
more options for producing good quality
micrographs. Adjusting the physical param-
eters of the SEM, such as, accelerating volt-
age, stigmator, focus, and spot size can now
be done with the touch of a button; adjust-
ing features of the image like contrast and
brightness is also made simpler. Most of the
newer SEM’s have the ability to digitize
images and save them as files for importa-
tion into image-enhancing software. Once
stored as a digitized image the user can then
opt for “‘cleaning-up”’ or enhancing image
quality after the fact.
Although technology is moving ahead on
the hardware end, the fact remains that
preparation of the specimen is still the most
important first step and this is the area
where little comparative work on procedu-
res has been conducted. Some new tech-
niques reported in the literature involve
lengthy processing, the use of chemicals or
materials that are difficult to handle, or spe-
cialized equipment not commonly available
(Grodowitz et al. 1982; Colwell and Kokko
1985). While these expenditures of time
and resources may be useful for intensive
study of a paticular taxon, the applicability
of these methods to other insect taxa or to
specimens of different body types is limit-
ed.
Our comparative analysis of methods
commonly used for SEM preparation
among different types of insects is a first
VOLUME 99, NUMBER 3
9696 100.80
Fig. 2.
Freezing fixation. A, Dialeurodes citri fourth instar nymph, dorsal view. B, D. citri, detail of dorsal
integumental reticulation. C, Bemisia argentifolii fourth instar nymph, dorsal view. D, B. argentifolii, detail of
the ventral marginal opening of the tracheal furrow. E, Encarsia sp., female, dorsal view of the thoracic region.
FE. sp., female, antenna.
step in determining the applicability of
these methods in obtaining consistent, qual-
ity specimens.
The procedure which ranked the highest
among all four taxa tested was ethanol fix-
ation, followed by osmium tetroxide post-
fixation, dehydration in ethanol, and criti-
cal-point drying. This procedure provided
generally clean specimens with little col-
lapse or charging. Its limitation may be the
loss of secreted surface features, such as
wax, thus, rendering the specimen “‘unlife-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
15KU X72 9916 100.80
oe
Be? ies. au
Fig. 3.
7
skKU X6600 © BOB7
15K 1300 if. @U
6611
Ethanol fixation. A, Dialeurodes citri fourth instar nymph, dorsal view. B, D. citri, detail of dorsal
integumental reticulation. C, Bemisia argentifolii fourth instar nymph, dorsal view. D, B. argentifolii, detail of
the ventral marginal opening of the tracheal furrow. E, Encarsia sp., female, dorsal view of the thoracic region.
FE. sp., female, detail of a funicle segment of the antenna.
like’, but again, better suited to viewing
integumental surface features. This method
has also been used successfully to examine
a wide array of insect taxa and insect stages
(DHH, unpublished data). Ethanol fixation
has produced the best and most consistent
results for SEM examination of insect eggs
(DHH, unpublished data). In the present
study, fixation by freezing also gave good
results and eliminated the need for the extra
rehydration steps before postfixing, as is
needed with ethanol fixation. Procedures 3
VOLUME 99, NUMBER 3
1S5KY #2006 BAG 1
Fig. 4. Ethanol fixation, HMDS, Encarsia sp., A,
Female, dorsal view of the thoracic region. B, Female,
detail of a funicle segment of the antenna. C, The apex
of the antennal club.
and 4 gave excellent results, but were not
widely applicable. Dipteran larvae are un-
usally prone to shrinkage and distortion
during processing for SEM. Further, they
are often contaminated with debris and oth-
er by-products of their various micro-habi-
Fig. 5. Hot-water fixation, HMDS, Musca domes-
tica. A, Gnathocephalon, right lateral view. B, Anterior
prothoracic spiracles. C, Terminal sensory organ of the
anterior sensory lobe.
tats that are not easily removed by chemical
means, e.g., baths in hexanes, bleach, etc.
Other taxa such as lepidopteran or coleop-
teran larvae can withstand such harsh treat-
ments without damage. We have found that
procedure 4 works exceptionally well to
22
provide clean, undistorted larvae; larger-
bodied forms are cut apart to facillitate pen-
etration during the fixation process.
The procedure that may be considered
the best will vary with the ultimate goal of
the user. For example, life-like appearance
may not be compatible with ethanol fixa-
tion. More taxa and body types must be
studied in a comparative fashion. The in-
clusion of preparation methods in the lit-
erature, however briefly, would aid devel-
opment of procedures that predictably pro-
vide clean, life-like specimens and good
quality SEM images.
ACKNOWLEDGMENTS
We thank Carol Meisenbacher and Bob-
bie Orr for technical assistance and the use
of specimens from their colonies. We also
thank R. D. Goeden and J. M. Heraty for
reviews of early drafts of this manuscript.
LITERATURE CITED
Adams, J. L., C. J. Battjes, and D. A. Buthala. 1987.
Biological specimen preparation for SEM by a
method other than critical point drying, pp. 956—
957. In Bailey, G. W., ed., Proceedings of the 45"
Annual Meeting of the Electron Microscopy So-
ciety of America. San Francisco Press, San Fran-
cisco.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Barbosa, P. 1974. Manual of Basic Techniques in In-
sect Histology. Autumn Publisher, Mass. 253 pp.
Belcari, A. 1987. Contributi alla concoscenza dei Dit-
teri Tefritidi. I[V. Descrizione della larva di terza
eta’ de Acanthiophilus helianthi (Rossi), Dacus
oleae (Gmel.), e considerazioni preliminari sulle
differenziazioni morfologiche legate al diverso
trofismo. Frustula Entomolgica 10: 83-1235.
Colwell, D. D. and E. G. Kokko. 1985. Preparation
of dipteran larvae for scanning electron micros-
copy using a freeze-substitution technique. Cana-
dian Journal of Zoology 64: 797-799.
Dawes, C. J. 1988. Introduction to Biological Electron
Microscopy: Theory and Techniques pp. 247-249.
Ladd Research. Industries, Inc., Burlington, Ver-
mont.
Gordh, G. and C. J. Hall. 1979. A critical point drier
used as a method of mounting insects from alco-
hol. Entomological News 90: 57-59.
Grodowitz, M. J., J. Krchma, and A. B. Broce. 1982.
A method for preparing soft bodied larval diptera
for scanning electron microscopy. Journal of the
Kansas Entomology Society 55: 751-753.
Humason, G. L. 1967. Animal tissue techniques. 2"
edition. W. H. Freeman and Company: San Fran-
cisco. 569 p.
Nation, J. L. 1983. A new method using hexamethyld-
isilane for preparation of soft insect tissues for
scanning electron microscopy. Stain Technology
58: 347-351.
Sabatini, D. D., K Bensch, and R. J. Barnett. 1963.
The preservation of cellular ultrastructure and en-
zymatic activity by aldehyde fixation. Journal of
Cellular Biology 17: 19-58.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 523-536
DESCRIPTIONS OF A NEW GENUS AND SIX NEW SPECIES OF NEARCTIC
LESTREMIINAE (DIPTERA: CECIDOMYITIDAE)
MATHIAS JASCHHOF
Zoologisches Institut und Museum der Universitat Greifswald, Bachstrasse 11/12,
D 17489 Greifswald, Germany.
Abstract.—A new genus and six new species of Nearctic gall midges are described as
new to science. These are: Amedia floridana, the only lestremiine totally lacking the
medial vein; Allarete bicornuta, Neurolyga longipes and Neurolyga pritchardi, all three
characterized by their remarkable male genitalia; Heterogenella californica, the first rec-
ord of this genus in North America and the first species within the tribe Bryomyiini
showing brachypterous females; and Polyardis occulta, representing a further case of
brachyptery in the female within the genus.
Key Words:
species
The lestremiine gall midges in the Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C., con-
tain a large number of unidentified or only
generically determined specimens, mostly
collected since the revisional work by Prit-
chard (1947a, 1947b, 1951). I studied this
collection for a revision in progress of the
Holarctic Lestremiinae. It contained six re-
markable species described here as new to
science. There is no further connection be-
tween these species, besides the fact that the
material was sufficient enough in number of
specimens and condition to describe the
species with confidence. The most striking
species, Amedia floridana, for which I erect
a new genus, lacks the medial vein com-
pletely. Allarete bicornuta possibly sup-
ports the view that Allarete in its present
composition is not a natural group of spe-
cies. Two new species of Neurolyga, lon-
gipes and pritchardi, represent two mor-
phological extremes among the species in-
cluded in this genus. Heterogenella califor-
nica, the first record of this genus in North
Diptera, Cecidomyiidae, Lestremiinae, Nearctic Region, new genus, new
America, is the first partially brachypterous
species described within the tribe Bry-
omyiini. Polyardis occulta is brachypterous
in the female and the third such case within
Polyardis.
Amedia Jaschhof, new genus
Type-species: Amedia floridana Jasch-
hof, new species, by present designation.
Adult male (female unknown).—Wings
without media; R; reaching wing margin
near wing apex; CuA unforked; CuP short;
macrotrichia on both sides of R;; antennae
with 14 flagellomeres, setae and sensoria of
flagellomeres not in regular whorls; dorsal
transverse bridge of gonocoxites with dis-
tinct apodemes, tergum 10 free, not fused
with tergite 9.
Remarks.—Apart from the lacking the
media, Amedia best fits the diagnostic char-
acters of the Strobliellini, where it is placed.
In the monotypic genera previously belong-
ing to this tribe, Strobliella Kieffer and
Groveriella Mamaev, M,,, is unforked and
obsolete distally, and a long M,,, is present.
524
Compared with the forked M,,, of the Ca-
totrichini, Lestremiini and Catochini, this is
an apomorphic character, as it is the com-
plete loss of the medial vein in Amedia. It
should be emphasized that this loss is not
combined with a general reduction of wing
size, veins and macrotrichia, that accom-
panies brachyptery in some _ lestremiine
genera. I hesitate to erect a unique tribe for
Amedia, because it is known only from a
single species and only the male. In addi-
tion, our knowledge of the species-poor
Strobliellini is insufficient as well. With the
exception of the reduction of medial veins,
Amedia is characterized by a combination
of plesiomorphic character states. The fla-
gellum resembles that of Catotricha species
(tribe Catotrichini) in lacking regular
whorls of setae or sensoria. The male ter-
minalia, with the comb-like structure at the
apex of the gonostylus and the trapezoidal
tergite 9, reminds one of some species of
Catochini.
Etymology.—The name Amedia is of
feminine gender and means “without me-
dia”’ referring to the unique wing venation.
Amedia floridana Jaschhof, new species
(Figs. 1-8)
Male.—Body size: 2,5—3,0 mm. Head:
High but short, appearing flattened frontal-
ly. Postfrons bilobed, slightly prominent,
with a few setae. Postcranium with sparse,
scattered, long setae and no postocular bris-
tles. Distinct suture present posterior to eye
bridge and reaching occipital foramen, ver-
tex on both sides of suture prominent. Clyp-
eus not developed, region between insertion
of antennae and mouth-parts flat and bare.
Ocelli lacking. Compound eyes small; eye
bridge short, strongly restricted and without
facets at vertex. Antenna: scape of same
size as pedicel; with 14 flagellomeres, each
successively longer; fourth flagellomere
(Fig. 8) with very short neck, node long,
cylindrical, with irregularly scattered short
setae and inconspicuous sensory hairs, fully
pubescent. Palpus (Fig. 6) 2-segmented,
segment 1 stemmed, with short sensory
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
hairs ventrally and a few setae laterally,
segment 2 slenderer and mostly longer than
first regardless of the stem, with a few short
setae. Thorax: Scutum with 2 lateral and 2
dorsocentral sparse rows of long setae; scu-
tum with smooth transition into scutellum.
Tarsomeres without scale-like setae. Claws
very slightly bent, with minute teeth. Em-
podia reduced to a few hairs. Halteres with
very long and narrow stem, latter with
sparse setae, knob with a row of setae in
prolongation of stem. Wings (Fig. 7) long
and narrow; Sc extending beyond level of
rs; C without distinct break near wing apex;
R, 6-7 times as long as rs; M completely
lacking; CuA strong, bent distally; CuP
short, close to CuA; membrane with a few
macrotrichia peripherally; macrotrichia
sparsely scattered dorsally on R, R,, R; and
ventrally on R,;. Pattern of sensory pores:
R, 1-2, rs 1 (sometimes on R; proximally),
R, 2-3 distally. Anal area with straight mar-
gin. Abdomen: Tergites and sternites with
long, sparsely scattered setae concentrated
along margins. Terminalia: gonocoxites
(Fig. 3) with short setae ventrally, fused
only in proximal fifth by a membraneous
link, dorsal transverse bridge with long
proximolateral apodemes; gonostylus (Fig.
1) long, slightly arched, tapering to tip (no-
ticeable from above, Fig. 2), with a comb-
like structure apically and about 5 incon-
spicuous ventral spines subapically; genital
rod (Fig. 4) simple, narrow and sclerotized;
tegmen (Fig. 4) narrow and parallel-sided,
parameral apodemes swept ventrally, with
basal transverse bridge, membranous on
apical third and with many short hyaline
“‘spines”’; tergite 9 (Fig. 5) plate-like, near-
ly trapezoidal, distal margin straight or
slightly emarginate, with short, scattered se-
tae; tergum 10 free, large, bilobed, with
long apical setae and covered with setulae;
sternum 10 about half length of tergum 10,
bilobed, with long apical setae and covered
with setulae.
Female.—unknown.
Types.—Holotype ¢, I-18-1964, swept
from Medicago sativa, Gainesville, Florida,
VOLUME 99, NUMBER 3
Pes
Maaco
+
3
yt
“"
z
“*
v
%
8
6
Figs. 1-8. Amedia floridana, male. 1, Gonostylus. 2, Gonostylus (detail of tip, from above). 3, Genitalia
(partial, left side: ventral, right side: dorsal). 4, Tegmen and genital rod (ventral). 5, Tergite 9 and tergum 10
and sternum 10 (dorsal). 6, Palpus. 7, Wing. 8, Fourth antennal flagellomere.
526
U.S.A., EW. Mead, deposited in National
Museum of Natural History, Washington,
D.C. Paratypes: 8 ¢, same data as holotype;
1 36, gopher tortoise burrow, undated, Hol-
lister, Putnam Co., Florida, U.S.A., E.G.
Milstray.
Remarks.—This unique species stands
out by the combination of characters dis-
cussed under the generic description. It is
further remarkable by its frontally flattened
head, the lack of ocelli, and the reduced
mouthparts and wing macrotrichia.
Etymology.—The name floridana refers
to Florida, the type locality.
Allarete bicornuta Jaschhof, new species
(Figs. 9-17)
Male.—Body size: 2.5—2.7 mm. Head:
Postfrons without setae. Eye bridge short
and 4 facets long, a little constricted at ver-
tex. Postcranium densely covered with
short setae. Scape markedly larger than
pedicel. Antenna with 14 flagellomeres;
first flagellomere on one side covered with
short sensory hairs along the whole length
of node; neck of fourth flagellomere (Fig.
17) slightly shorter than node; node with 1
basal whorl of setae, mesally 1 complete
crenulate whorl of extremely long setae and
2-3 short crenulate rows of long setae, dis-
tally many short sensory hairs and 1-2
large sensory spines; distal flagellomeres
with gradually fewer and shorter sensory
hairs. Palpus (Fig. 16) 4-segmented; seg-
ments increasingly longer distally; first seg-
ment with short sensory hairs; all segments
with setae. Thorax: Scutum with 2 lateral
and 2 dorsocentral rows of long setae. Tar-
someres without scale-like setae. Claws
very short and strong, slightly arcuate; ba-
sally with several teeth, each successively
longer. Empodia reduced to some hairs.
Halter with setae on basal half. Wing (Fig.
14): h present; Sc not reaching level of rs;
R, as long as in other Allarete species; rs
and r-m very short; M-fork markedly lon-
ger than stem of M,,,; M, broader and
darker than M,, both veins parallel-sided,
diverging only distally; CuA slightly sinu-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ous distally; CuP present; A long, apparent
only in central section. Macrotrichia present
on all veins with exception of h, rs, r-m and
M; macrotrichia on both sides of R;, M,,>
(distally) and M-fork (proximally). Pattern
of sensory pores: R, 2(—3), R; 2 proximally,
3 distally. Anal lobe strongly convex. Ab-
domen: Tergites and sternites with long se-
tae. Terminalia: gonocoxites (Fig. 9) with
relatively short setae ventrally except on
basal third and membraneous medial link,
with ventral lobes, dorsal gonocoxal apo-
demes strong and connected by a weak
transverse bridge; gonostylus (Figs. 9, 10)
long, broadest at base, tapering to tip, the
distal fourth arched inwards, with finger-
nail-like apical tooth and 4—5 short spines
subapically; genital rod (Fig. 12) sclero-
tized, with membraneous pubescent cap
apically, 2 inconspicuous acute projections
from apex of genital rod directed proxi-
molaterally; tegmen (Fig. 12) membra-
neous, parameral apodemes sclerotized and
directed ventrally, centrally with an y-sha-
ped sclerotization, both distal sides of y
with double-toothed apex; tergite 9 (Fig.
11) wide, nearly membraneous, densely
scattered with setae, distal margin straight;
tergum 10 large, free, bilobed, with long
and strong setae dorsally and apically; ster-
num 10 obviously onelobed, with rounded
distal margin and covered with setulae.
Female.—Body size: 3.0—4.0 mm. Head:
Antenna: scape little smaller than pedicel;
with 9 flagellomers, the last constricted and
with small second node; first flagellomere
with 2 small depressions with short sensory
hairs (such sensory depressions slightly no-
ticeable also on second to seventh flagel-
lomeres); fourth flagellomere (Fig. 13) bar-
rel-shaped, with very short neck, 1 whorl
of long setae in basal third and many short
sensory hairs. Thorax: Tarsomeres of fore-
legs with many short, spine-like setae on
the ventral surface (found distally on first
tarsomere) and indistinct on tarsomeres of
other legs. Abdomen: With 1 large, flat-
tened, poorly sclerotized spermatheca (ir-
VOLUME 99, NUMBER 3 527
16
Figs. 9-17. Allarete bicornuta. 9, Male terminalia (partial, left side: ventral, right side: dorsal). 10, Gon-
ostylus (detail of tip, dorsal). 11, Tergite 9 (dorsal). 12, Tegmen and genital rod (ventral). 13, Female fourth
antennal flagellomere. 14, Male wing (partial). 15, Spermatheca. 16, Male palpus. 17, Male fourth antennal
flagellomere.
528
regular margin possibly resulting from de-
formity, Fig. 15).
Types.—Holotype 4, light trap, VII-16-
1958, Cochise Stronghold, Dragoon Moun-
tains, Arizona, U.S.A., C.W. O’Brien, de-
posited in National Museum of Natural His-
tory, Washington, D.C. Paratypes: | d and
5 2, same data as holotype.
Remarks.—Allarete was established by
Pritchard (1951) for those Lestremiini with
a medial fork with branches both being of
same vein width. This character is of little
value, since within all species included in
Allarete M, is little broader and darker than
M, (even if not so clearly indicated as in
some Lestremia or Anaretella species).
Probably Allarete is not a natural group of
species, but, in the present sense includes
species with these plesiomorphic wing
characters: the long R, and A; macrotrichia
on both sides of R;, M,,, including fork and
A; and 2 (not 1) sensory pores on R,; prox-
imally. We are not able at present to place
all Lestremiini with confidence into natural
groups. Therefore the new species is re-
ferred to Allarete with reservation and for
lack of alternatives. The structure of the
tegmen is unusual compared with the other
congeners and within the tribe, as is the oc-
curence of a spermatheca.
Etymology.—The name bicornuta is an
adjective and means “‘with two horns”’ and
refers to the distal projections of the teg-
men.
Neurolyga longipes Jaschhof,
new species
(Figs. 18-24)
Male.—Body size: 3.5 mm. Head: Post-
frons without setae. Eye bridge with a few
scattered facets laterally, medial eye portion
to 4 facets long. Occiput sparsely covered
with setae posteriorly. Postgenae with long
setae, | inconspicuous row of 4—5 postoc-
ular bristles. Scape clearly larger than ped-
icel. Antenna with 12 flagellomeres; neck
of fourth flagellomere (Fig. 24) clearly lon-
ger than node; node with 2 irregular whorls
of setae basally, 1 complete and 3 incom-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
plete crenulate whorls of long setae (distal
flagellomeres with 2 complete crenulate
whorls) and groups of short sensory hairs
distally. Palpus (Fig. 23) long, 4-segment-
ed, segments gradually increasing in length,
first segment with short sensory hairs in-
side, all segments with setae. Thorax: Scu-
tum with 2 lateral and 2 dorsocentral sparse
rows of long setae. Legs extremely long,
clearly longer than body. Claws arched at
right angle, distal side longer than proxi-
mal, toothed. Empodia longer than claws.
Halteres densely covered with short setae.
Wings longer than body; Sc slightly reach-
ing level of rs; rs short, R, 10—11 times as
long as rs; M,,, apparent also distally;
CuA, not reaching wing margin; CuP pres-
ent, nearly %4 as long as CuA,,,. Macro-
trichia on membrane and on R, R,, R; and
r-m (only distally). Pattern of sensory
pores: R, 3—4, rs 1, R; 1 proximally, 1-3
medially/distally. Anal lobe clearly convex.
Abdomen: Tergites sparsely covered with
long setae, more concentrated laterally.
Sternites densely covered with long setae.
Terminalia: gonocoxites (Fig. 18) with a
widened base without setae ventrally, oth-
erwise with long setae except a broad stripe
along medial line, distal margin with u-
shaped neckline, inner bridge of gonocoxi-
tes with remarkably long setae; gonostylus
(Fig. 19) long, slightly arched inwards,
rounded distally, with long setae very
densely covering distal third; genital rod
(Fig. 20) strongly sclerotized, apically wid-
ened to membraneous head with minute
hairs, mouth of sperm ducts apparently
leading into head of genital rod proximo-
laterally; tegmen (Fig. 21) with long and
sclerotized parameral apodemes, nearly par-
allel-sided, separated along two thirds of
medial line, distolaterally with 2 character-
istic angled and acute projections directed
laterally; tergite 9 (Fig. 22) plate-like,
slightly sclerotized, distal margin broadly
rounded, distally with bilobed projection
with strong, stiff, inwardly directed hair;
tergum 10 fused with tergite 9, bilobed and
covered with setulae.
VOLUME 99, NUMBER 3 529
Figs. 18-24. Neurolyga longipes, male. 18, Terminalia (partial, left side: ventral, right side: dorsal). 19,
Gonostylus (dorsal). 20, Tip of genital rod. 21, Tegmen (ventral). 22, Tergite 9 and tergum 10 (dorsal). 23,
Palpus. 24, Fourth antennal flagellomere (anterior).
530
Female.—unknown.
Types.—Holotype <4, sticky trap, II-25-
1976, Willamette National Forest, Oregon,
U.S.A., Voegtlin & Christy, deposited in
National Museum of Natural History,
Washington, D.C. Paratype: 1 ¢, same data
as holotype.
Remarks.—This new species is unique
among the Neurolyga (formerly Cordylo-
myia, see below) in that the body size of
the adults is much larger in relation to other
species of this genus. The legs, antennae,
palpi and wings are long in proportion to
body size. The occurence of CuP and a dis-
tinctly convex anal lobe are plesiomorphic
conditions usually not appearing within the
Micromyidi. The male genitalia support the
view that Neurolyga and Campylomyza
form a monophyletic group. They share the
medially divided tegmen with sclerotized
projections, and the over all shape of the
aedeagus corresponds as well. This species
is referred to Neurolyga because of the ab-
sence of setae on the katepisternite and the
lack of angled macrotrichia on wing mem-
brane, the presence of both being synapo-
morphies of Campylomyza.
Remark on nomenclature.—I studied the
remains of the original material of Neuro-
lyga fenestralis Rondani, 1840, the type
species of Neurolyga, in the Rondani col-
lection. As a result, I consider N. fenestralis
to be identical with Cordylomyia coprophi-
la Felt, 1911, the type species of Cordylo-
myia. Consequently, Neurolyga is no longer
considered a junior subjective synonym of
Campylomyza Meigen, 1818, as proposed
by former authors, and Cordylomyia Felt,
1911, is a junior subjective synonym for
Neurolyga Rondani, 1840.
Etymology.—The name longipes is a
noun in apposition meaning long legs.
Neurolyga pritchardi Jaschhof,
new species
(Figs. 25—29)
Male.—Body size: 1.4—1.7 mm. Head:
Postfrons without setae. Eye bridge 1 sparse
row of facets long laterally, medial eye por-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tion to 3—4 facets wide. Occiput sparsely
covered with setae posteriorly; postocellar
bristles indistinct. Postgenae sparsely cov-
ered with long setae, with a short row of
about 3 postocular bristles. Antenna with 12
flagellomeres; neck of fourth flagellomere
(Fig. 29) little shorter than node; node with
1 whorl of basal setae, 1 complete and 3
incomplete crenulate whorls of long to very
long setae and groups of short sensory hairs
distally. Palpus 4-segmented; last segment
longest; first segment with short sensory
hairs inside, very few sensory hairs also on
second and third segments; all segments
with setae. Thorax: Scutum with 2 lateral
and 2 dorsocentral sparse rows of long se-
tae. Tarsomeres without scales. Claws
arched at right angle and toothed. Empodia
reaching % to % of claw length. Halter
densely covered with short setae. Wing: Sc
not reaching level of rs; R, 3 times as long
as rs; M,,, very faint; CuA, short, not
reaching wing margin. Macrotrichia cover-
ing membrane and on R, R,, r-m, R; and
CuA,,>. Pattern of sensory pores: R, 3, rs
1, r-m 1, R; 1 medially/distally. Abdomen:
Tergites with a few setae dorsally, more
densely covered with setae laterally. Ster-
nites with long setae. Terminalia: gonocox-
ites (Fig. 25) covered with long setae ven-
trally with exception of a stripe along the
medial line, distal margin with narrow
u-shaped neckline extending to more than
¥% of gonocoxite length, dorsal transverse
bridge with long apodemes proximolater-
ally; gonostylus (Fig. 26) arched inwards in
distal third, with fingernail-like tooth api-
cally surrounded by 4 long inconspicuous
spines; genital rod (Fig. 27) long and
strongly sclerotized, widened and forked
apically, tips of fork sclerotized; tegmen
(Fig. 27) broadest at base, parameral apo-
demes clearly swept ventrally, on both sides
with narrow acute projection directed ven-
trally, distally 2 notched projections obvi-
ously seperated by a gap; tergite 9 (Fig. 28)
large, plate-like, distal margin broadly
rounded, with long scattered setae; tergum
10 fused with tergum 9, bilobed, densely
VOLUME 99, NUMBER 3 S3il
Figs. 25-29. Neurolyga pritchardi, male. 25, Terminalia (partial, left side: ventral, right side: dorsal). 26,
Gonostylus (ventral). 27, Tegmen (ventral). 28, Tergite 9 and tergum 10 and sternum 10 (dorsal). 29, Fourth
antennal flagellomere.
532
covered with setulae and a few setae; ster-
num 10 inconspicuous and covered with
short setulae.
Female:—unknown.
Types.—Holotype ¢, IV-27-1947, Inver-
ness, California, U.S.A., A.E. Pritchard, de-
posited in National Museum of Natural His-
tory, Washington, D.C. Paratype: 1 4,
IV-27-1947, Lagunitas, California, U.S.A.,
A.E. Pritchard.
Remarks.—Within the genus the new
species is unusual for the apical tooth-like
structure of the gonostyli. Only one other
Neurolyga species has that character: UN.
subbifida (Mamaev 1963) described from
European Russia. I was not able to obtain
material for comparative study and the de-
scription of subbifida is unsufficient. The
sketchy available figure showing terminalia
(Mamaev 1963: 452, Fig. 4 g) indicates that
both species are similar in the apex of teg-
men and genital rod as well. The tegmen of
N. subbifida is described as having two di-
verging teeth and two further teeth of gen-
ital rod between them. This is also the case
in N. pritchardi, but the tegmen of N. prit-
chardi has additional lateral acute projec-
tions directed ventrally. Recently I was in-
formed by B. M. Mamaev, that WN. prit-
chardi is not identical with N. subbifida, a
decision being founded on a comparison of
subbifida with my drawings of pritchardi.
The new species is separated from the con-
geners by the short empodium and the pat-
tern of sensory pores on wing veins: usually
Neurolyga species have 3 pores on R,, | on
rs, | on R; proximally and 2 medially/dis-
tally. The pattern of pores in subbifida was
not described.
Etymology.—The new species is named
to honor A. E. Pritchard, who collected the
type material, for his outstanding contribu-
tions to the taxonomy of Nearctic lestre-
miines.
Heterogenella californica Jaschhof,
new species
(Figs. 30-35)
Male.—Body size: 1.2—-1.8 mm. Head:
Postfrons without setae. Eye bridge 2-3
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
facets wide. Postcranium - except a stripe
behind eye bridge - with long setae, post-
genae also with scales. Antenna with 12 fla-
gellomeres; neck of fourth flagellomere
(Fig. 34) little to clearly shorter than node;
node narrow, with | basal whorl of setae, 1
complete and 3 incomplete crenulate whorls
of long setae and 5—6 sensory hairs of var-
ious length distally (2—4 of the them long
and strong, mostly 1 bi- or three-furcated).
Palpus 4-segmented; fourth segment lon-
gest; first segment with short sensory hairs
inside; all segments with setae and scales.
Thorax: Scutum with 2 dorsocentral and 2
lateral sparse rows of short setae. Tarso-
meres with setae and broad scales. Claws
crescent-shaped, without teeth. Empodia
only just as long as the claws. Halter cov-
ered with narrow scales. Wing: Sc not
reaching level of rs; R, 2.5—3.5 times as
long as rs; M,,, obsolete distally; CuA-fork
very acute; CuA, not reaching wing mar-
gin; membrane densely covered with ma-
crotrichia and sparsely on R, R, proximally
and CuA,,,. Only 1 (not 2 as usual) sensory
pore on R; medially/distally. Abdomen: Ter-
gites with sparse row of setae dorsally and
small patch of setae laterally. Sternites with
long setae and scales. Terminalia: gonocox-
ites (Fig. 30) with long setae ventrally, dis-
tal margin with broadly u-shaped neckline,
without projection on both sides of neck;
gonostylus (Fig. 31) widest in distal half,
clearly excavated on inside distally (notice-
able only in lateral view), distal third dor-
soventrally flattened and remarkably dense-
ly and strongly covered with setae, apex
rounded; the base of the genital rod (Fig.
32) sclerotized, otherwise membraneous
and bifurcated distally; tegmen (Fig. 32)
membraneous, more or less parallel-sided,
distal margin broadly rounded; tergite 9
(Fig. 33) plate-like and nearly trapezoidal
(proximal margin seems to be excavated,
but, this results from deformity by cover
glass pressure), distolaterally 2 pointed
lobes covered with setulae and directed in-
wards; tergum 10 fused with tergite 9 and
bilobed, densely covered with long setulae;
VOLUME 99, NUMBER 3 2333)
Ln Ni Fe
(yy / a .
—~L fad } H
se +
Se es / “A sae
e ‘\ ee ;
A
cr (= Ip oe ay nS “A
; ee! 4
aS , 3 ds.
x ™, ye" a
g y aes :
Figs. 30-35. Heterogenella californica. 30, Male terminalia (partial, left side: ventral, right side: dorsal).
31, Gonostylus (dorsal). 32, Tegmen and genital rod (ventral). 33, Tergite 9 and tergum 10 and sternum 10
(dorsal). 34, Male fourth antennal flagellomere. 35, Female fourth antennal flagellomere.
sternum 10 large, bilobed, densely covered _ tered facets long laterally and without facets
with strong setulae. at the vertex. Antenna with 8 flagellomeres,
Female.—Head: Eye bridge shorter than _ last flagellomer constricted mesially; fourth
in male; in brachypterous form 1-2 scat- flagellomere (Fig. 35) slender, bottle-
534
shaped, with long neck; node with | sparse
whorl of short sensory hairs basally, 1
whorl of long setae above that, | row of
sensory hairs mesally and two 2- to
5-forked sensoria distally. Palpus 3- or
4-segmented. Thorax: Macropterous and
brachypterous (to nearly apterous) forms
occur side by side. Thorax of brachypterous
form stunted, mesonotum very sparsely
covered with short setae. Stubs of wing as
long as thorax or clearly shorter, partly with
indistinct venation and some macrotrichia
at margin. Tarsomeres also of brachypter-
ous form with broad scales. Abdomen: Seg-
ments with short setae. 2 small, poorly
sclerotized, rounded and flattened sperma-
thecae.
Types.—Holotype 6, IV-27-1947, Inver-
ness, California, U.S.A., A.E. Pritchard, de-
posited in National Museum of Natural His-
tory, Washington, D.C. Paratypes: 5 d and
5 2, from redwood litter, III- 18-1953, Oak-
land, California, U.S.A., W.C. Bentinck.
Remarks.—This is the first record of Het-
erogenella in the Nearctic Region, and H.
californica is the only known species of the
genus with wing reduction. It is remarkable
that brachypterous and fully winged fe-
males occur simultaneously. The only other
known female of Heterogenella is that of
H. bigibbata Mamaev and Berest, resem-
bling H. californica in all respects with the
exception of the wing reduction. The male
terminalia of H. californica are character-
ized by gonostyli widened and excavated
inside in distal half and with a dense tuft of
long setae apically. All other Heterogenella
males exhibit a uniform covering with setae
of gonostyli.
Etymology.—The name californica re-
fers to California, the type locality.
Polyardis occulta Jaschhof, new species
(Figs. 36—43)
Male.—Body length: 1.1 mm. Head:
Postfrons with | strong seta. Eye bridge 1—
2 facets long laterally, 2—3 facets at the ver-
tex. Occiput with a few strong postocellar
bristles, without setae elsewhere. Postgenae
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
sparsely covered with long setae. Antenna
with 13-15 flagellomeres; neck of fourth
flagellomere (Fig. 40) as long as the node
or little shorter; node with 1 whorl of long
basal setae, 1 complete and 1 incomplete,
poorly developed crenulate whorls of long
setae mesally and further setae and sensory
hairs of various length distally. Palpus (Fig.
41) 2-segmented; first segment enlarged,
globular, densely covered with short sen-
sory hairs inside; second segment pointed,
shorter and narrower than the first; both
segments with a few setae. Thorax: Scutum
with 2 dorsocentral and 2 lateral sparse
rows of short setae. Coxae, femura and tib-
iae of all legs very strong; tarsomeres with-
out scales. Claws crescent-shaped; untooth-
ed. Empodia as long as claws. Halter
sparsely covered with short setae. Wing: Sc
not reaching level of rs; R, 1,5 as long as
rs; CuA, strong, but short; wing membrane
without macrotrichia, R, R, and R; with a
few macrotrichia only. Abdomen: Tergites
with a few setae only laterally. Sternites
densely covered with long setae. Termina-
lia: gonocoxites (Fig. 36) with long to very
long setae ventrally, distal margin with
broadly v-shaped neckline; gonostyli (Fig.
37) small, broadest in proximal third, sharp-
ly tapering to tip, with 1 long, slender api-
cal tooth and 2 inconspicuous ventral spines
subapically; genital rod (Fig. 39) slender,
opening in distai fifth, somewhat widened
(ovoid) and lightly sclerotized apically; teg-
men (Fig. 39) shield-shaped, with rounded
apex; tergite 9 (Fig. 38) with narrow scler-
otized posterior margin and scattered setae;
tergum 10 fused with tergite 9, bilobed,
densely covered with setulae and with a few
fine setae; sternum 10 hidden, bilobed and
covered with setulae.
Female.—Body size: 1.2 mm. Head: Eye
bridge 1—2 rows of scattered facets long.
Postcranium sparsely covered with setae.
Antenna with 9 flagellomeres; first flagel-
lomer with field of many short sensorial
hairs on prolonged basis; fourth flagellomer
(Fig. 42) tapering in distal half, with incon-
spicuous neck; node with a few sensory
Nn
oS)
Nn
VOLUME 99, NUMBER 3
Figs. 36-43. Polyardis occulta. 36, Male terminalia (partial, left side: ventral, right side: dorsal). 37, Gon-
ostylus (dorsal). 38, Tergite 9 and tergum 10 and sternum 10 (dorsal). 39, Tegmen and genital rod (ventral). 40,
Male fourth antennal flagellomere. 41, Male palpus. 42, Female fourth antennal flagellomere. 43, Spermatheca.
536
hairs and 1 whorl of setae basally and many
short sensory hairs apicad of the setae. First
segment of palpi stronger enlarged than in
male. Thorax: Stunted in size; mesonotum
with a few short setae only. Brachypterous;
wings noticeable as short stubs without ve-
nation. Fifth tarsomere of forelegs twice as
long as fourth. Abdomen: Setae very short.
1 large, sclerotized, disc-shaped spermathe-
ca (Fig. 43), on one side with many small
pores.
Types.—Holotype d, shrubby meadow,
VII-15 to VIII-28-1977, 18 km east of Gan-
anogue, Ontario, Canada, Dondale & Red-
ner, deposited in National Museum of Nat-
ural History, Washington, D.C. Paratypes:
2 3 and 2 2, same data as holotype.
Remarks.—The species represents the
third case of brachyptery in the female
within Polyardis beside P. silvalis (Ron-
dani) and P. recondita (Lengersdorf) (Jas-
chhof, in prep.). Like other wing reduced
species in different genera, P. occulta ex-
hibits characters frequently connected with
brachyptery or aptery beside the remark-
able reduction of the thorax. These are the
reduced vestiture on the head and meso-
notum, the reduced number of facets of the
eye bridge, shortened palpi and partially
broadened legs. The winged male is partly
affected as well, not only by the shortened
palpi, but, also by a reduced vestiture with
macrotrichia on the wing and the poorly de-
veloped crenulate whorls on the flagello-
meres. The new species is the only Polyar-
dis species with 2-segmented palpi, an ex-
ceptional character within the whole sub-
family Lestremiinae. The male genitalia are
of the usual type and similar to the conge-
ners, but the gonostyli are small and sharply
tapered with a longer and slenderer apical
tooth compared to other species. In addi-
tion, the genital rod opens in the distal fifth
and exhibits an ovoid widening apically.
Etymology.—The name occulta is an ad-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
jective and means occult, referring to the
combination of exceptional characters, es-
pecially in female sex.
Remark on nomenclature.—Polyardis
Pritchard, 1947 is not a junior subjective
synonym of Campyloneura Lengersdorf,
1939, as I previously considered it to be
(Jaschhof and Menzel 1995). Recently I
was informed by Mr. K. Voigt, Ettlingen,
that Campyloneura is a preoccupied name
(by Campyloneura Fieber, 1860, in Miri-
dae, Heteroptera).
ACKNOWLEDGMENTS
I am very grateful to R. J. Gagné, Sys-
tematic Entomology Laboratory, USDA, %
National Museum of Natural History,
Washington, D.C., for reading and discuss-
ing the manuscript with me, for his helpful
criticisms, and for the loan of the specimens
used in this study. I thank a unknown re-
viewer of the manuscript for his corrections
and comments. A study trip to investigate
the collections of the National Museum of
Natural History, Washington, D.C., in No-
vember, 1994 was supported by the Deut-
sche Forschungsgemeinschaft (DFG).
LITERATURE CITED
Jaschhof, M. and E Menzel. 1995. Campyloneura
Lengersdorf, 1939—eine “‘vergessene”’ Lestremi-
inen-Gattung wiederentdeckt (Diptera, Cecido-
mylidae, Lestremiinae). Beitrage zur Entomologie
45: 427-430.
Mamaev, B. M. 1963. Gallicy fauny SSSR, triba Mi-
cromyini Rondani. Entomologicheskoe Obozrenie
42: 437—454. (In Russian)
Pritchard, A. E. 1947a. The North American Gall
Midges of the tribe Micromyini; Itonididae (Ce-
cidomyiidae); Diptera. Entomologica Americana
(new series) 27: 1—87.
. 1947b. The North American Gall Midges of
the tribes Catotrichini and Catochini (Diptera:
Itonididae (Cecidomyiidae)). Annals of the Ento-
mological Society of America 40: 662-671.
. 1951. The North American Gall Midges of
the tribe Lestremiini; Itonididae (Cecidomyiidae);
Diptera. University of California Publications in
Entomology 8: 239-275.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 537-540
MORPHOLOGICAL CHARACTERISTICS OF THE STING AND PREY
CARRIAGE MECHANISM IN SERICOPHORUS RELUCENS F. SMITH
(HYMENOPTERA: SPHECIDAE: LARRINAE)
IvicA T. RADOVIC AND SRDJAN SUSIC
Institute of Zoology, Faculty of Biology, University of Belgrade, Studentski trg 16,
Belgrade, Serbia.
Abstract.—Morphological characteristics of the sting concerning prey carriage in Ser-
icophorus relucens F Smith are described. The sting apparatus consists of sting palpi
(with bristles), stylet, and lancets, with different morphological adaptations. Correlation
between the morphological features of some parts of the sting and their function in prey
carriage are discussed.
Key Words:
There are basically three types of prey
carriage in the Sphecidae: mandibular,
pedal, and abdominal, each with several
subtypes. The most primitive one is man-
dibular (found in the majority of Spheci-
dae species), and the more advanced are
pedal (found in four groups of this family)
and abdominal (found in three groups of
this family) (Evans 1962). The abdominal
type of prey transport includes three sub-
types: abdominal transport with the use of
a specially modified apical abdominal
segment, “‘buprestid clamp’’ subtype, a
modification of the fifth abdominal ster-
num, and abdominal transport with the
use of the sting.
The first subtype is found in Listropy-
gia and Clypeadon (Philanthinae), and is
characterized by a modified apical abdom-
inal segment in the form of a clamp
(Evans 1962; Bohart and Menke 1976).
The second subtype is found in certain
Cerceris species (Krombein 1981).
The third subtype is found in some Cra-
broninae (Oxybelus sp., Crossocerus sp.)
(Evans 1962) and in Sericophorus relu-
cens F Smith (Larrinae) (McCorquodale
Sericophorus relucens F. Smith, Sphecidae, prey carriage, sting morphology
1988). McCorquodale (1988) first report-
ed sting prey carriage in a species outside
the Crabroninae, S. relucens, the first
found in Larrinae which is one of the most
advanced prey transport mechanisms.
Most of the Sphecidae that use their
sting apparatus for prey transport exhibit
some adaptive changes in the morphology
of several structures of the sting, mainly
the stylet, that are correlated to scleroti-
sation and mobility of their prey (Radovic
1985). Those changes in general are a
curved stylet in sphecids that prey on fast
flying insects; a slightly curved or straight
stylet in those praying on slower insects.
The presence of bristles on the distal part
of palpi and presence of barbs on the dis-
tal part of lancets function in securing
prey impaled on the stylet.
Sericophorus relucens preys upon Dip-
tera (Matthews and Evans 1970; Peckham
and Hook 1980). They sting their prey in
the air, on the ventral side of the thorax,
and keep it impaled on the sting, not re-
moving it while flying, till they get to the
nest (McCorquodale 1988).
538 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1—4.
the palpus (SEM 100%) (original). 3, Tip of the stylet (SEM, 100%) (original). 4, Lancet barbs positioned in
lancet tracks (SEM, 100%) (original).
MATERIALS AND METHODS
We examined five specimens obtained
from Dr. O. Lomholdt (Zoologisk Museum,
Copenhagen), from Blue Mountains and
Kurrajong, New South Wales; Mount
Wedge and Mundiwindi, Western Australia.
The sting apparatuses (Fig. 1) were first
removed from the abdomen of the speci-
mens, and initially preserved in glycerol.
They were cleaned for scanning electron
microscopy (SEM) by soaking in 1:1 solu-
tion of ammonium hydroxide and water, de-
hydrated in ethanol, soaked in chloroform,
and air dried. We fixed them to the SEM
stubs using silver conducting paint. A
Sericophorus relucens. 1, Lateral view of the sting apparatus (SEM, 65%) (original). 2, Tip of
“sputter coater’’ was used for gold coating
the objects (Gibson 1984). The stings were
filmed using “‘Foma”’ (5 X 5 cm) film.
RESULTS
Examining five specimens of S. relucens,
we found the following elements of the
sting apparatus:
1. Sting palpi.—Two sting palpi consist-
ing of two segments are present (Fig. 1).
Several (commonly around ten) bristles are
found on the tip of the distal part (Fig. 2).
Bristles are distributed along the horizontal
axis of the palpus tip forming two groups:
longer bristles (on the distal part of the tip)
VOLUME 99, NUMBER 3
and shorter bristles (on the sides of the tip).
The bristles are unsegmented, rough sur-
faced, straight with slightly enlarged base.
Similar bristles are found on the side of the
proximal segment of the palpi.
2. Stylet——The stylet (Fig. 3) is long
and curved (less than in Oxybelus species)
with a sharp pointed tip. The surface and
edges of the stylet are rough, covered with
tubercles, and different in shape and size.
The lancet tracks, concavely shaped with no
special structures (like olistheter scales or
setae found in Vespidae, Ondricek-Fal-
Ischeer 1992), are located on the ventral
side of the stylet, and covered with larger
tubercles. The cracklike poison duct is lo-
cated near the tip of the stylet.
3. Lancets.—A pair of lancets is located
on the ventral side of the stylet. We found
unexserted lancets in all of the examined
specimens. Lancets are sharply pointed and
covered with small tubercles. Similar to
some crabronine sphecids, the lancets of S.
relucens are barbed (Fig. 4). Six barbs are
positioned on the dorsal side and form a
line parallel to the longitudinal axis of the
lancet. The barbs differ in size: those closer
to the lancet’s tip have smaller dimensions
in contrast to those with proximal position
that are particularly wider at their base. No
variation was found in the number of barbs
between the two lancets and among differ-
ent specimens. Sensilla campaniformia
were not noted.
DISCUSSION
Regarding morphological and behavioral
characteristics, the type of prey carriage
found in S. relucens belongs to the abdom-
inal ‘‘sting’”’ type that is found only in two
other genera of Sphecidae and is considered
to be one of the most advanced (Evans
1962).
Most of the features of the sting appa-
ratus examined in our research show cor-
relation between their morphological char-
acteristics and their function in prey car-
riage, generally in securing contact of the
sting with the body of the victim.
559
Sting palpi are provided with receptory
bristles that can also function in holding the
body of the prey. The majority of sphecids
that prey upon fast fliers (Diptera, Hyme-
noptera) have a curved stylet (Radovic
1985). The stylet of S. relucens, although it
preys upon Diptera, is not extremely curved
(compared to that in Oxybelus species),
probably due to light prey preference. Nu-
merous tubercles on the sides and edges of
the stylet function in increasing the stylet
surface that is in contact with the victim’s
internal tissues. As the lancet barbs are po-
sitioned dorsally, the sharp pointed stylet
tip probably does not function in severing
parts of the victim’s tissues caught in the
lancet barbs.
The function of lancets is anchoring the
stylet in the body of the victim (Ondricek-
Fallscheer 1992). We found no lancets in
exserted position in all specimens exam-
ined, so we could not ascertain whether the
lancets protrude beyond the tip of the stylet.
Dorsally positioned barbs have a function
in securing prey impaled on the sting (Ra-
dovic 1985).
Most of the listed structural modifica-
tions are correlated to complex behavior
patterns of this species. Predatory insects
that use their sting for prey carriage share
several important advantages to those that
use mandibles or legs. Positioning the prey
in the back, wasps shift their gravitational
center that is important in maintaining over-
all balance; this type of carriage permits
rapid provisioning of the nest that positive-
ly affects the prey searching distance, pro-
vides fewer opportunities for predators and
parasites to attack the prey, and leaves the
mandibles and legs free for use otherwise
(after Evans 1962).
CONCLUSIONS
There are three basic types of prey car-
riage in the family Sphecidae: mandibular,
pedal and abdominal, each with several
subtypes. The abdominal type of prey trans-
port includes three subtypes one of which
540
is abdominal transport with the use of the
sting, such as that found in S. relucens.
Using SEM we found the following in S.
relucens: two sting palpi consisting of two
segments with approximately ten receptory
bristles distributed along the palpus tip that
also function in securing prey impaled on
the sting and the long and curved stylet
with a sharp pointed tip, covered with tu-
bercles which are different in shape and
size, and provided with two lancet tracks;
lancets, positioned on the ventral side of the
stylet, and embedded in lancet tracks, are
sharply pointed and covered with small tu-
bercles, six barbs are found on the dorsal
side of the lancets.
Most of the structural characteristics ex-
amined serve in securing firm contact of the
sting with the body of the victim.
ACKNOWLEDGMENTS
This research was supported by the Ser-
bian Republic Fund for Science, project No.
03E04. We are also grateful to Dr. Ole
Lomholdt for specimens used in this re-
search.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
LITERATURE CITED
Bohart, R. M. and A. S. Menke. 1976. Sphecid Wasps
of the World, a generic revision. University of
California Press. Berkeley, 695 pp.
Evans, H. E. 1962. The evolution of prey-carrying
mechanisms in wasps. Evolution 16(4): 468—483.
Gibson, G. 1984. Specimen preparation and scanning
electron microscopy. Sphecos 9: 8-10.
Krombein, K. V. 1981. Biosystematic Studies of Cey-
lonese Wasps, VIII: A Monograph of the Philan-
thidae (Hymenoptera: Sphecoidea). Smithsonian
Contributions to Zoology. No. 343.
Matthews, R. W. and H. E. Evans. 1970. Biological
notes on two species of Sericophorus from Aus-
tralia (Hymenoptera: Sphecidae), Psyche 77: 413—
429.
McCorquodale, D. B. 1988. Prey carriage on the sting
by Sericophorus relucens (Hymenoptera: Spheci-
dae: Larrinae). Journal of New York Entomolog-
ical Society 96(1): 121-122.
Ondricek-Fallscheer, R. L. 1992. A morphological
comparison of the sting apparatuses of socially
parasitic and nonparasitic species of yellowjackets
(Hymenoptera: Vespidae). Sociobiology 20(3):
245-300.
Peckham, D. J. and Hook A. W. 1980. Behavioral
observations on Oxybelus in southeastern North
America. Annals of the Entomological Society of
America 73: 557-567.
Radovic¢, I. T. 1985. Morphology and adaptive value
of the sting apparatus of digger wasps (Hym.,
Sphecidae). Acta Entomologica Jugoslavica 21(1—
2): 61-73.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 541-559
BIOLOGY OF FOUR SPECIES OF NOTIPHILA FALLEN (DIPTERA:
EPHYDRIDAE) ASSOCIATED WITH THE YELLOW WATER LILY,
NUPHAR LUTEUM (NYMPHAEACEAE)
LEN LARSON AND B. A. FOOTE
Department of Biological Sciences, Kent State University, Kent, OH 44242, U.S.A.;
(LL) current address: Department of Natural Sciences, Shawnee State University, Ports-
mouth, OH 45662, U.S.A.
Abstract:—The life histories of four species of Notiphila, N. bella, N. eleomyia, N.
kentensis, and N. nudipes, that feed on anaerobic sediments surrounding roots of the
yellow waterlily, Nuphar luteum, are described. Males of N. eleomyia and N. nudipes
form mating aggregations resembling leks on flower buds, flowers, fruits, and occasionally
leaves of the host plant, and mating occurs within these aggregations. Females deposit
large clusters of eggs on these plant parts, particularly the flowers and fruits. The incu-
bation period lasts 4—6 days, and newly hatched larvae drop off the egg masses and fall
into the underlying sediments where they feed during the summer and fall months. Over-
wintering occurs as nearly mature larvae. In late May and early June, larvae move to
roots of Nuphar and insert their spiracular spines into air spaces within the root tissue.
The pupal period lasts 14—22 days. Notiphila bella is univoltine, but the three other species
may be bivoltine. Information is presented on the utilization of wetland plants by different
species of Notiphila, and it is proposed that speciation and adaptive radiation involved
movement onto these different “‘host plants” for the purpose of obtaining oxygen by the
larvae.
Key Words:
Wetlands subjected to long-term inunda-
tion develop anaerobic conditions in the un-
derlying soils (Mitsch and Gosselink 1992).
As a result, plants and animals living in
such anaerobic conditions must acquire a
variety of mechanisms that allow them to
acquire the oxygen necessary for their met-
abolic functions. Many wetland plants pos-
sess aerenchyma in their stems, rhizomes,
and roots that allows for the diffusion or
mass transport of oxygen from the photo-
synthesizing aerial portions of the plant to
the roots (Armstrong 1964, 1978; Dacey
1980, 1981; Grosse and Mevi-Schutz 1987;
Mevi-Schutz and Grobe 1988). Larvae of
some insects living in anaerobic soils have
Shore flies, Notiphila, Ephydridae, Diptera, Nuphar, Ohio
modified posterior spiracles that form elon-
gate spines that can be inserted into the
aerenchymatous tissues of plants and thus
acquire oxygen (Miiller 1922; Varley 1937,
1939; Keilin 1944; Hinton 1953. 1968;
Hartley 1958; Houlihan 1969a, 1969b). A
large and widely distributed taxon having
this ability are shore flies belonging to the
genus Notiphila Fallén.
The genus Notiphila is worldwide in dis-
tribution, occurs in a variety of wetland
habitats, and is speciose, with 53 species
having been recorded from America north
of Mexico (Mathis 1979, Wirth et al. 1987).
Meaningful biological studies are available
for only five of these Nearctic species, al-
542
though fragmentary information has been
published for an additional 15. Berg (1950)
reared N. (Notiphila) loewi Cresson from
anaerobic soils surrounding the rhizomes
and roots of pondweeds (Potamogeton
spp.). Eastin and Foote (1971) discussed the
natural history of N. (Dichaeta) caudata
Fallén, a species that is unusual within the
genus in that its larvae are not buried in
anaerobic sediments but crawl about on the
surface. Larvae of this species lack spirac-
ular spines. Busacca and Foote (1978) pre-
sented life history observations on two spe-
cies, N. (Agrolimna) aenigma Cresson, and
N. (N.) solita Walker, that obtain oxygen
from roots of cattails (Typha latifibula L.).
They also described the eggs, larvae, and
puparia of these two species. Deonier et al.
(1978) described the immature stages and
outlined the natural history of N. (N.) car-
inata Loew, a species that is associated with
water willow (Justicia americana (L.)
Vahl.) growing in slow-flowing streams in
southwestern Ohio and other regions east of
the Mississippi River.
Some biological observations and indi-
cations of association with particular wet-
land plants are also available for N. (N.)
bella Loew, N. (N.) eleomyia Mathis, N.
(N.) kentensis Huryn, N. (N.) mathisi Hu-
ryn, N. (N.) nudipes Cresson, N. (N.) pau-
roura Mathis, N. (N.) poliosoma Mathis, N.
(N.) shewelli Mathis, N. (N.) taenia Mathis,
N. (N.) theonae Huryn, and N. (A.) deonieri
Mathis, N. (A.) olivacea Cresson, N. (A.)
quadrisetosa Thomson, WN. (A.) scalaris
Loew and WN. (A.) scoliochaeta Mathis
(Mathis 1979; Todd and Foote 1987; Huryn
1984, 1987).
In this paper, we present observations of
the natural history of Notiphila bella, N.
eleomyia, N. kentensis, and N. nudipes oc-
curring in stands of the yellow water lily,
Nuphar leuteum (L.) Sibth. and J. E. Smith.
We also discuss resource partitioning in the
genus Notiphila and propose a possible
mode of speciation within this taxon.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
MATERIALS AND METHODS
Site description.—Two marshes located
in Portage County in northeastern Ohio
(Fig. 1) were sampled in this study. The
larger marsh is situated on the east side of
State Highway 43, 9.3 km north of the
northern city limits of Kent. The marsh is
bordered on the west by the highway, res-
idential property on the north, and open
lowland woodlands on the east and south.
The emergent marsh vegetation consists of
species of Carex, Typha, and Sparganium
around the perimeter, a large stand of Typha
and Sparganium growing intermixed on a
floating mat of dead vegetation at the east
end, and a large stand (ca. 2700 m7’) of yel-
low water lily at the west end (Fig. 2). In
addition to the emergent vegetation, there
are thick growths of floating duckweeds,
Lemna minor L., Wolffia punctata Griseb.,
Spirodela polyrhiza (L.) Schleid., and water
milfoil, Myriophyllum exalbescens Fernald.
The marsh is persistent and has a rather sta-
ble water level. The water depth varied
from 50 to 80 cm during the two-year study
period.
The second, much smaller marsh is lo-
cated 0.8 km east of the Kent State Uni-
versity main campus, between Horning
Road and U.S. Highway 261 (Fig. 3). It is
bordered by Horning Road on the west, res-
idential property and a pond on the north,
Highway 261 on the east, and an abandoned
agricultural field on the south. The vegeta-
tion is more diverse than in the larger marsh
and consists of discontinuous stands of Ty-
pha, Carex, Phalaris, Sparganium, Eleo-
charis, and Nuphar. At the time this marsh
was characterized by Todd and Foote
(1987), the Nuphar stand encompassed
some 1157 m’, but low water levels since
that survey has permitted Typha to invade.
As a result, the continuous stand of Nuphar
in 1995 is only some 100 m?’, and the water
depth averages only 10 cm.
Sampling methods for adults.——A tran-
sect was established across the center of the
Nuphar stand at the Route 43 marsh (Fig.
VOLUME 99, NUMBER 3 543
Fig. 1. Map showing locations of two study marshes in northeastern Ohio.
544
WOODED
2
WOODED
MARSH #2
Figs. 2-3.
showing stands of Nuphar luteum.
2). The transect extended south-southeast
from the north edge of the marsh. A rope,
65 m long, that was attached at each end of
the transect completely crossed the width of
the marsh. The Nuphar stand along the
transect was 45 m wide. Six sampling sites
were established along the transect at 6 m
intervals. Each site was marked by wooden
stakes that encompassed a 4 X 2 m plot.
Two sampling sites were established in the
Horning Road marsh (Fig. 3). One of these
was in the center of the Nuphar stand, the
other some 5 m distant where Nuphar and
Typha were intermixed.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
LAWN
58
TYPHA/SPARGANIUM
on Moating mat of
WOODED
dead TYPHA leaves
MARSII #1
Center pan trap
Edge pan trap
2, Map of the Route 43 marsh showing stands of vegetation. 3, Map of Horning Road marsh
Collections of adult Notiphila were taken
weekly at 2 locations in each marsh be-
tween early June and early October from
1991 to 1993. All collections were made
during the same 24 hour period using iden-
tical detergent pan traps (Southwood,
1978). The pan traps consisted of white
polyethylene storage containers measuring
30 X 15 X 10 cm. Each trap contained ap-
proximately 50 ml of water into which one
ml of dishwashing detergent had been add-
ed. All traps were placed in the marshes
during the mid-afternoon and collected 24
hours later. In the Route 43 marsh, traps
VOLUME 99, NUMBER 3
Figs. 4—5.
were placed at sampling sites 1 and 3,
whereas in the Horning Road marsh, they
were placed at both collecting sites. The
contents of the traps were returned to the
laboratory where the adult Notiphila were
removed, identified, sexed, and counted.
Collections of adult Notiphila from the
unopened buds or flowers of the Nuphar
were obtained either by using a modified
battery-operated hand vacuum (Marshall
1982) or by carefully placing a large poly-
ethylene bag over the buds or flowers and
then breaking off the peduncle below the
water surface and sealing the bag.
During the summer of 1992, six emer-
gence traps were placed along the transect
in the Route 43 marsh, one trap at each end
of the sample plots. The traps were con-
structed by attaching a one-piece plastic
tube pan to the outside bottom of a plastic
3-gallon (11.25 L) bucket (Fig. 4). Prior to
attaching the pan, a small hole was cut in
the center of the bucket. A lid of clear,
polycarbonate plastic was attached to the
bucket with machine screws and wing nuts.
A 5 cm vent hole was cut in the center of
the lid and covered with plastic window
screening. The two holes allowed for vent-
3.5 CM SQUARE WOOD
POLE 2.2 M LONG
8 MM THICK X 50 MM
avrres DIAMETER RUBBER PAD
3 MM THICK X 40 MM DIAMETER
POLYCARBONATE DISK
50 MM GALVANIZED 5
ROOFING NAIL
4, Emergence trap, cross-section diagram. 5, Diagram of core sampling device.
ing but prevented rainwater from flooding
the trap. Traps were attached to the wooden
stakes delimiting the sample plots and the
bucket part of each trap was completely
submerged. Each tube pan contained ap-
proximately 500 ml of water and one ml of
detergent. The traps were emptied weekly
between June 15 and July 27, 192, and the
Notiphia identified, sexed, and counted.
Sampling methods for larvae.—Core
samples of the bottom substrates were taken
at the sample sites in the Route 43 marsh
on September 20, 1991 (site 1), February
18, September 25, October 8, 1992, and
April 6, 1993 (all six sites). Cores were ob-
tained using a 5 cm colorless, polybuterate
tube that was 2 m long (Fig. 5). A piston
fitted into the coring tube allowed the sub-
strate to be sampled without being flooded
by the overlying water. Cores were placed
in polyethylene bags and returned to the
laboratory where they were examined for
Notphila larvae. Larvae were extracted
from the cores either by using an elutriator
similar to that described by Whitman et al.
(1983) or adding water to the mud sample
to reduce it to a slurry that was then spread
546
over the bottom of a large tray and inspect-
ed visually.
Rearing methods.—Portions of Nuphar
rhizomes and attached roots were removed
from the marsh substrate at various times
throughout the year and examined visually
in the laboratory for attached larvae and/or
puparia. Puparia were carefully removed
from the roots and placed on moist sphag-
num moss in test tubes (Berg 1950). They
were oriented in the moss so that the pos-
terior spiracular spines of the puparia were
exposed to air. No attempt was made to rear
larvae.
Identification of adults —Males were de-
termined by using keys of Mathis (1979)
and Huryn (1987). Tentatively determined
specimens of each species were submitted
to Wayne Mathis at the Smithsonian Insti-
tution for confirmation.
RESULTS OF TRAPPING IN THE Two
MARSHES
Pan trapping in the Route 43 marsh.—
Approximately 53,200 adult Notiphila of
eight species (Table 1, Fig. 6) were ob-
tained in the detergent pan traps during the
three-year survey. Of these, 10,620 (20.4%)
were males. Three species, N. eleomyia, N.
kentesis, and N. nudipes dominated
throughout the study (Figs. 7—9), represent-
ing 95.2% of the total number collected.
These three, plus N. bella, constituted over
97% of the males collected. N. eleomyia
was the most abundant species, accounting
for 60.3% of the total number. The second
most abundant species was N. kentensis, ac-
counting for 25.5% of the males collected,
whereas N. nudipes accounted for only
9.4% of the males identified. Notiphila bel-
la was present only during late spring and
early summer and commonly was the first
species to emerge (Fig. 10). Four other spe-
cies, N. olivacea, N. pauroura Mathis, N.
solita, and N. taenia, were only collected
sporadically. Notiphila taenia was taken
predominantly from mid to late June, al-
though a few were taken as late as October.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Temporal distribution of adults of all spe-
cies of Notiphila coilected in two marshes in Portage
County, Ohio.
Horning
Route 43 marsh Road
Week ee emia
number 1991 1992 1993 199]
19 5 * = *
20 * * ES *
21 s * 41 *
22 * * 544 *
73, 2 1451 3898 %
24 * 4309 10,410 *
25 2850 2052 8864 254
26 1925 1370 3524 79
27 1056 3316 ° 66
28 381 as * 4
29 233 2322 * 2
30 131 617 : 3
31 122 337 = —
32 440 160 | =
33 219 174 ce =
34 * 95 Kk *
35 66 372 « —
36 58 338 e =
37 50 * * -
38 8 131 * ==
39 2 67 * =
40 24 * x ies.
4] * * ** *
42 * 116 * *
Totals 7565 17,386 27,274 408
* No collections made.
The other three species were taken irregu-
larly throughout the summer months.
Forty-nine percent of the adults were
taken in pan traps located near the center of
the Nuphar stand (site 3), and 51% from
the edge of the stand (site 1) (Fig. 11). The
only species that did not follow this distri-
butional pattern was N. pauroura, as only
25% of its adults were taken in the center-
most traps.
The species composition of Notiphila re-
mained fairly stable over the three years,
but abundances changed considerably. The
total number of individuals collected in
1991 was 7565, but this increased every
year to the 27,275 trapped in 1993 (Fig.
12). The maximum number of flies trapped
over a 24-hour period (Fig. 12) occurred
VOLUME 99, NUMBER 3
100005-
NUMBER OF FLIES
8
eleomyia kentensis nudipes
\ ESS
XS NS ~~
WM
solita
NOTIPHILA SPECIES
Fig. 6.
within days of the same date on all three
years, VI-19-91, VI-17-92, and VI-16-93.
Pan trapping in the Horning Road
marsh.—Only 408 adults (38.2% males) of
five species of Notiphila were taken in the
Horning Rd. marsh during the year-long
survey (Table 1). The relative abundances
of the five species were somewhat different
than those obtained in the larger marsh. No-
tiphila eleomyia and N. kentensis accounted
for 51.9% and 20.5%, respectively, of the
adults trapped. However, the third most
abundant species was N. pauroura, ac-
counting for 13.5%, followed by N. nudipes
(11.5%) and N. olivacea (2.6%).
Emergence traps.—Somewhat surpris-
ingly, only nine males and 15 females of
three species, N. eleomyia (5), N. nudipes
(3), and N. bella (1), were collected in the
Total number of Notiphila males collected in the Route 43 marsh during 1992.
emergence traps. All appeared between
VI-17 and VI-29-92. Most of the adults
were taken at site 1, and none was taken in
the trap at site 4.
Core sampling.—Twenty Notiphila lar-
vae, all third instar, were extracted from the
core samples of mud that were taken inter-
mittently in the marshes (Table 2). No lar-
vae where reared to the adult stage, so spe-
cies identification was not possible.
Rhizome and root sampling.—A total of
229 larvae and 91 puparia were collected
from samples of Nuphar rhizomes and roots
(Table 3). All of the larvae were found dur-
ing the late fall and early spring months in
mud adhering to the rhizomes and roots. In
contrast, all puparia were firmly affixed to
the primary roots of the rhizomes and roots
(Fig. 21) and were found only during late
548
MALE N. ELEOMYIA
MALE N. KENTENSIS
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
800
\
700 cweccwcecesrnnscscseseecnesseccececccescosesesssecesescecssesecsccsess N |e neveeeeereeereeereecseececcccccccccessccececesessssessceeeeecesssseseseeeessessse een eneeeeeesseeseerreseneserenraseasreseeseeneneranamaseenereeseenessussasascnsuaes!
nh
Nw
0 ee N- N [ea i ete Nn aan i meme or la ae ac at eee SEM A ae
Nw
NaN
Nw
500 NO A a ert ‘a N Nec pocseboaceaceccscueesscncsascecacecceces cooreovseeesecceoceeseceeccscscusscscccacesnsosesesunnsanabanvvenccseessnnasennaldseedéesunsennews=denanusesuasnesanaaane|
\) : s)
Ni Nay
0 0 i ee yw... A el lee sean 2. eB ee Sees St Sere ee ee
Ney ON
Ny y N
) y N
500 eae N N- yoo
‘NB Ss
y : N
DOO porns rerecceeeeeeeettttetrnttrenenecnencnnnn 7 = N~ ve N
yh NRW
yn AS
100 Deen PaaS se re eee Sere eset sane ocescy N” No NN NN Fee ee psec croc nee ea em ee cet es nb se ee ete es ce eee
YR NQW\ N
N i NNWNNNN gf XN :
19 20 21 22 23 24 25 26 27 28 299 WH 32 BS 3 BS 37 38 39 40
WEEK OF THE YEAR
450
400 on enn nacancanaaddonnbansavanescerecrsbesses esesesseal tp cascecarsbeseonssestonssesseusssacoucerescpasvesa sees sosasaese os savvsbepsceceess pvorbeve==seuersrsuusrsns q=nur=snnansanne=Spee~e ==nse>sseeres
BBO enn ceceectneeenenenenenenenehenenenenenentnnnenenenenenenntntntnnmneetnannntnenenanenantreneneatnensernenenesnetnty
300 a a ee ee) Ee. ee cee
DBO sovrnrnncrccececeeernenemenenenene hace ereflneeeeneneneeenennenentnenennenntnennantanstansntnantenentntntnenesnetenenmnmnnanannnees
200 See 8 SS eee
150 en enn by ae - Se ee cccosllll uc norescevedenseseteteterincenanss-ssesteeeddnscosenes Ssass=aassdusetssapasiamesas tanssppep se ses a Sotrsn ret eEOCery
\
\
100 Lai an a en ne emery men encones | be a fe Nae Be ae ee
\ N
\ \ N
50 occencnenavcasnsoases=coeneabeverssneesvonscssnsoepsene = yoo \eg yp) SS === SS
‘ a)
sD SWS. MS
19 20 21 22 23 24 25 26 27 28 29 30 31 33 34 35 36 37 38 39 40
WEEK OF THE YEAR
1991 J 1992 1993 8
Figs. 7-8. 7, Seasonal distribution of Notiphila eleomyia in the Route 43 marsh. 8, Seasonal distribution of
N. kentensis in Route 43 marsh.
VOLUME 99, NUMBER 3 549
200
1 60 ensnvaccncacsrccccccsscccccvcscsceserereessnenccncccscsceseceses Sh ponnced Goccccccccccssccsccsccccsscccessecsnsncccscsecscneessoesatererssscecseccees-seeeneersrsnccenesececececeacacccccecsscececceseceececaceacececese:
”
Lu
s
Q Y DO savrrnrrerrerrrrttertrterttettttetecnncnneecceedbceeneecsnennsnnnnneeeennnennneesssenesnnnnnnsnnsssoneseesseneeneeeemessnanenerenamneeseaneaeeeeeeeeeeceeseseceeceeeeeenetesteeneeueeeetees
>
ZS
Z|
uw 80 aad peatnemn sere eeneee coin inenon een eranatnrensnaneesanedceadcemses Boece mae Soe Cn RE a re Se
—
6
= .
29) al kan ena eee eee IN ‘ cS eS es ee ee
NBay
N NR
NBS SN
mn i &) NS “ir : : :
19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 35 36 37 38 39 40
WEEK OF THE YEAR
IY 1991 BE 1992 Wy 1993 :
. BELLA
Z|
MALE
5 26 27 28 29 30 31 32 33 35 36 37 38 39 40
WEEK OF THE YEAR
AW 1991 BB 1992 1993
Figs. 9-10. 9, Seasonal distribution of adult Notiphila nudipes in the Route 43 marsh. 10, Seasonal distri-
bution of adult Notiphila bella in the Route 43 marsh.
10
550
NUMBER OF FLIES
Figs. 11-12.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
eleomyia kentensis nudipes bella pauroura taenia olivacea solita
SPECIES OF NOTIPHILA
EDGE |g CENTER
Peak Is the maximum number of Notiphila collected In a 24 hour 27275 (05/26-06/30)
period, both pan traps.
NUMBER OF FLIES
(Thousands)
12
of Notiphila collected in the Route 43 marsh between 1991 and 1993.
11, Spatial distribution of adult Notiphila in the Route 43 marsh during 1992. 12, Total number
VOLUME 99, NUMBER 3
Table 2. Larvae collected in core samples taken at
the Route 43 marsh in Portage County, Ohio.
Date Site 1 Site2 Site3 Site4 Site5 Site 6
09/20/91 9 ° eS os eS zt
02/18/92 3 0) 10) 0) 0) 0)
09/25/92 2 0 1 0) 0) 10)
10/08/92 1 0 1 0 0) 0
04/06/93 a 0 0) 10) 1 0
Totals 17 (0) 2 0) 1 (0)
* Not sampled.
May and early June. Frequently, several pu-
paria were attached to the same primary
root, and there appeared to be a preference
for the smaller roots located near the apical
end of the rhizome. Some 20 puparia found
on 10 roots of a single rhizome were all on
roots that measured from 1.1 to 2.8 mm in
diameter at the point where the root at-
tached to the rhizome.
ADULT BEHAVIOR
The adult behavior of the species studied
in this investigation resembled that de-
scribed for other species of Notiphila by
Busacca and Foote (1978), Deonier et al.
(1978), Simpson (1976) and Van der Velde
and Brock (1980). Adults spent most of
their time walking over the surfaces of the
Nuphar leaves and stems. They flew only a
few centimeters when disturbed. While on
the leaves, adults frequently paused to
‘‘taste’’ the surface, feeding on a microflora
or absorbing the abundant honeydew de-
posited by a large leafhopper (Homoptera:
Delphacidae) population. They were also
observed to feed on bird droppings.
Courtship and Mating Behavior.—The
flower buds, expanded flowers, and fruits of
Nuphar were sites of a courtship aggrega-
tion of males that resembled lekking (Hég-
lund and Alatalo, 1995). Large numbers of
males, usually of a single species, distrib-
uted themselves rather evenly over the
flower (Fig. 13) or bud (Fig. 14). They
faced downward towards the water surface
and held their wings out approximately 30
degrees from the centerline of the body,
Nn
Nn
—_
Table 3. Number of Notiphila taken from roots of
Nuphar luteum at the Route 43 marsh.
Date Site 2 Site 4 Site 6
09/20/91 Larvae 5 ay
Puparia O “
05/28/92 Larvae 1 1 i
Puparia 49 0) 14
06/03/92 Larvae 0) ** *
Puparia 21 zs *
04/06/93 Larvae 122 SY 4
Puparia 0) 0) 0)
04/27/93 Larvae 26 6 13
Puparia 3 O 0)
* Not sampled.
forming a v-shape (Fig. 15). Males con-
stantly jostled each other and maintained a
fairly uniform spacing. Whenever an inter-
loper landed on the flower, it was quickly
pounced upon by the nearby flies. If the
newcomer was a male, he joined the group
after a bit of shuffling to accommodate him.
If the visitor was a female, there was a
struggle among the males for her attention
until one individual was able to copulate
with her. Once the female was mounted, the
other males resumed their positions on the
flower head and ignored the mating pair.
Occasionally, a male attempted to disrupt
the mating pair, but these efforts were usu-
ally unsuccessful.
Males were particularly attracted to flow-
ers that possessed numerous Notiphila eggs.
Of male aggregations on 29 flowers, 16
flower buds, and two fruits that were ex-
amined, all but one possessed eggs (Fig.
16). On one occasion, an aggregation of six
males of N. eleomyia was observed on a
leaf that supported 775 eggs (Fig. 17).
Aggregations were observed in N. eleo-
myia and N. nudipes but not in N. bella, N.
kentensis, or N. taenia. Mating behavior in
those 3 species remains unknown. Male ag-
gregations of N. nudipes typically were on
the expanded flowers (418 of 488 males
collected on flowers were of this species),
whereas males of N. eleomyia were more
commonly found on the buds (119 of 223
os
NUMBER OF FLIES
g
100
NUMBER OF FLIES
Figs. 13-14.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
nudipes eleomyia taenla
SPECIES OF NOTIPHILA
MALE [ij FEMALE 13
Each column represents a single bud or frult
32 32 MMM BB BW 37 BF F7
WEEK NUMBER
N. nudipes [ij N. eleomyia 14
13, Total number of Notiphila on expanded flowers of Nuphar luteum at the Route 43 marsh
during 1992. 14, Number of males of Notiphila on flower buds of Nuphar luteum at the Route 43 marsh during
1992
N
1)
ie)
VOLUME 99, NUMBER 3
J) RP SEAS
i |
MARAARR ARORA AEA
Ae
Figs. 15-20. 15, Aggregation of males of Notiphila nudipes on flowers of Nuphar luteum. 16, Eggs of
Notiphila sp. on the underside of flower bud of Nuphar luteum. 17, Eggs of Notiphila eleomyia on the underside
of a leaf of Nuphar luteum. 18, Close-up of Notiphila eleomyia. 19, Spiracular spine of larva of Notiphila sp.
20, Puparium of Notiphila sp. attached to primary root of Nuphar.
554
males collected on buds were of this spe-
cies).
Mating.—Mating activity was observed
in two pairs of adults of N. eleomyia on two
flower buds that also supported several oth-
er males. Each flower bud also had 60—90
Notiphila eggs. In the first pair, mating last-
ed for six minutes. After initial copulation
was completed, the male continued to ride
the female for another six and a half min-
utes and then re-initiated copulation, re-
maining in copula for an additional two
minutes. The female terminated copulation
by pushing at the male with her hind legs.
She remained on the bud, whereas the male
flew away. In the second pair, copulation
lasted four and a half minutes. The male
then continued to ride the female for anoth-
er 12 minutes before decamping.
Copulating females frequently moved
about on the surface of the leaves while
feeding on the accumulated honeydew.
They commonly carried the male with them
as they moved to the flowers for oviposi-
tion.
Oviposition.—The egg masses consisted
of as few as six to a maximum of 615 eggs.
Most of the egg masses were deposited pri-
marily on the reproductive parts of Nuphar,
although a few were found on leaves. Less
than one percent of 300 leaves examined
weekly along the transsect possessed egg
clusters, whereas over 96% of the examined
flowers supported egg masses (Table 4). In
large egg masses on buds, flowers, and
fruits, eggs were deposited on all surfaces
of these reproductive parts and frequently
consisted of two or three layers. In contrast,
masses deposited on leaves were always on
the lower side of the leaf and usually con-
sisted of only one layer of eggs.
The number of eggs laid per female dur-
ing any one time interval varied from spe-
cies to species. Females seemingly matured
and deposited a full complement of eggs at
one time. Two field-collected females of N.
eleomyia deposited clusters of 25 and 27
eggs, respectively, over 24 hours in a breed-
ing jar, suggesting that 12-14 eggs were be-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 4. Number of leaves and flowers of Nuphar
luteum having eggs of Notiphila.
Number of | Number of Number of | Number of
Leaves Leaves Flowers Flowers
Week Examined With Eggs Examined With Eggs
24 300 0 Bs ee
PLS) 300 0) a ~
26 300 6 = =
27 300 | “s =
28 300 3 18 18
29 300 0) 6 6
30 300 i 10 9
31 e = 10 8
a2, aS = 10 10
33 as s 10 10
34 s 2 10 10
35 = 8 8
36 = = 7 6
* Not sampled.
ing released from each ovary. The ovaries
of two dissected gravid females of N. eleo-
myia each consisted of 14 ovarioles. Each
ovariole contained two fully formed eggs
plus several others that were still undergo-
ing oogenesis. In contrast, only six to eight
ovarioles were found in each ovary of five
dissected N. nudipes.
Eggs of both N. eleomyia and N. nudipes
are white in color, show distinct texturing
on the upper surface (Fig. 18), and are
smooth on their ventral side. A distinct line
separates the two areas. The textured dorsal
surface of the egg may serve as a plastron
that allows respiration to continue even
when eggs are immersed in water (Hinton
1953, 1968; Deonier et al., 1978). Van der
Weld and Brock (1980) demonstrated that
eggs of N. brunnipes Robineau-Desvoidy,
an European species, developed normally
and hatched when submerged in well-aer-
ated water. The incubation period for eggs
of N. eleomyia and N. nudipes was three to
four days. A longitudinal line of weakness
at the micropylar end of the egg split open
at the time of hatching, allowing the larva
to emerge.
The only observed enemy of the eggs
was an undetermined species of Tricho-
grammatidae (Hymenoptera). Every egg of
VOLUME 99, NUMBER 3
a cluster of 300 located on a Nuphar leaf
was parasitized by this wasp.
LARVAL BEHAVIOR
Newly hatched larvae dropped from the
egg masses and moved into the anaerobic
sediments surrounding the roots of Nuphar.
They fed on these sediments during the
summer and fall, usually attaining the third
instar before overwintering. Feeding re-
sumed the following spring, and larvae at-
tached themselves to the Nuphar roots in
May (Fig. 19). The puparia were firmly af-
fixed to the roots (Figs. 20, 21) throughout
the pupal period, which lasted 14—25 days.
Notiphila bella appeared to be univoltine,
and was the first species to emerge as adults
(third week of May). No adults of this spe-
cies were collected after the third week of
June. The remaining three species may be
multivoltine, although the data are incon-
clusive. One possible indication of a mul-
tivoltine habit was the discovery of eight
adults of N. eleomyia on October 8, 1992,
whose ptilina were still exposed. All of
these species emerged later (early June)
than N. bella and were present in the marsh-
es over a longer period, with adults of three
species being encountered as late as early
October. Notiphila populations peaked in
late June and steadily declined as summer
progressed.
DISCUSSION
There is little evidence that competition
was an important factor among the five spe-
cies of Notiphila that dominated the Nuphar
guild, and resource partitioning was not
overly apparent. One possible example of
partitioning was the fact that N. bella was
univoltine and emerged earlier than the oth-
er four species. The two species that formed
male aggregations preferred different sites,
as males of N. nudipes formed aggregations
on open flowers, whereas those of N. eleo-
myia aggregated on flower buds. Larvae of
all five species co-occurred within the an-
aerobic sediments and apparently fed on the
organic detritus. They all attached them-
555
selves to roots of Nuphar before forming
puparia.
The huge number of adults obtained in
the pan traps (Fig. 22) strongly suggests
that the various species of Notiphila com-
prising the Nuphar-associated guild expe-
rience ecological release and thus can attain
very large populations. We encountered
very few other insect larvae in the anaero-
bic sediments that could be competing for
the same food resource with Notiphila, and
predators were rarely encountered.
There is a strong correlation of the life
cycle of the Nuphar-associated species of
Notiphila with the time of maximum oxy-
gen richness in the roots of the water lily.
Movement of oxygen from the leaves to the
stems and into the roots involves diffusion
and mass transport of gases driven by pho-
tosynthesis occurring in the aerial portions
of Nuphar. Oxygenation of the rhizome and
associated roots is particularly apparent in
the spring when newly formed leaves are
beginning to appear (Dacey 1981). This en-
sures that the developing pupae have access
to a rich source of oxygen at a time when
demand is particularly high.
An interesting question concerning No-
tiphila is why the genus is so speciose? It
is the second largest genus of Ephydridae
in North America (53 spp.) being exceeded
only by the phytophagous genus Hydrellia
(60 spp.) (Wirth et al. 1987, Mathis and Za-
twarnicki 1995). There is a general feeling
among systematists that genera comprised
of plant-feeding taxa are more speciose
than genera composed of predators or scav-
engers (Mitter et al. 1988). Speciation in
phytophagous insects apparently occurred
as populations shifted to new host plants
and gradually adapted to the particular set
of environmental conditions presented by
the different host taxa. The large genus Hy-
drellia is composed of leaf-mining and
stem-boring species and thus follows the
expected pattern. On the other hand, species
of Notiphila are basically detritivores, feed-
ing on anaerobic sediments in wetland hab-
itats. What could have driven such prolific
556 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 21-22. 21, Respiratory spines of puparium of Notiphila sp. embedded in primary root of Nuphar
luteum. 22, Adults of Notiphila in detergent in water pan trap collected over a 24-hour period.
VOLUME 99, NUMBER 3
Table 5. Known ‘‘host plants’? of North American
species of Notiphila.
Notiphila Species Host Plant Genus Source
The adusta group
The pulchrifrons group
N. bella Nuphar This paper
N. kentensis Nuphar This paper
N. mathisi Nuphar Huryn 1984
N. nudipes Nuphar This paper
N. taenia Nuphar This paper
N. theonae Nuphar Huryn 1987
The Joewi group
N. carinata Justicia Deonier et al.
1978
N. eleomyia Nuphar This paper
N. loewi Potamogeton Berg 1950
N. pauroura Nuphar This paper
N. poliosoma Potamogeton Mathis 1979
N. shewelli Nuphar Mathis 1979
N. solita Typha Busacca &
Foote 1978
N. scoliochaeta Carex Mathis 1979
The bispinosa group
N. bispinosa Spartina Foote, pers.
obs.
N. furcata Spartina Foote, pers.
obs.
The scalaris group
N. aenigma Typha Busacca &
Foote 1978
N. decoris Glyceria Foote, pers.
obs.
N. macrochaeta **Grasses”’ Mathis 1979
N. olivacea Nuphar This paper
N. quadrisetosa Agrostis Mathis 1979
N. scalaris Leersia Foote, pers.
obs.
speciation in this genus? We suggest that
speciation occurred as populations within
species of Notiphila adapted to different
wetland plants as sources of oxygen. All
species of Notiphila, except for those of the
caudata group of the subgenus Dichaeta,
are thought to obtain their oxygen from
aerenchymatous tissue in the roots of wet-
land plants. As Table 5 shows, 22 species
of Notiphila occurring in North America
have known “host plants’, although the
plant associations of 31 species are still un-
55) f/
known. Generalizations about suggested
plant-insect relationships within the genus
thus must be very tentative. The plants uti-
lized by the five species of the avia group
are completely unknown, and only one of
the three species comprising the pulchrif-
rons group has a suspected host plant. One
generalization solidly supported by evi-
dence is that species of Monocotyledonae
are preferred over species of Dicotylondeae.
Only N. carinata, has been reported from a
species of dicot (Justicia americana (L.)
Vahl), whereas 21 species have been asso-
ciated with monocots. Another well-sup-
ported generalization is that the genus Nu-
phar is heavily used by Notiphila larvae. At
least 10 species of three different species
groups representing both subgenera are
known, or suspected, to attach their puparia
to roots of species of yellow water lily. The
adusta species group may be restricted to
Nuphar, as six of its 11 species are consid-
ered to be affiliated with yellow water lilies.
Finally, species of the furcata and scalaris
species groups seem to show a preference
for species of grasses, as six of the 19 spe-
cies in those groups are known to utilize
Gramineae. An interesting divergence from
the typical Notiphila feeding pattern is pos-
sessed by species of the caudata group, as
larvae of the only reared species, N. di-
chaeta, feed on surface sediments and do
not possess spiracular spines.
ACKNOWLEDGMENTS
Appreciation is expressed to the Scheffel
family of Streetsboro, Ohio, for granting
unrestricted access to the marsh on their
property. We also extend appreciation to W.
N. Mathis, Department of Entomology,
Smithsonian Institution, Washington, D.C.,
for his advice and aid in determining adult
Notiphila.
LITERATURE CITED
Armstrong, W. 1964. Oxygen diffusion from the roots
of some British bog plants. Nature 204: 801—802.
Armstrong, W. 1978. Root aeration in the wetland
condition, pp. 269-297. In Hook, D. E. and R. M.
558
Crawford, eds., Plant Life in Anaerobic Environ-
ments. Ann Arbor Science Publishers, Ann Arbor,
Michigan.
Berg, C. O. 1950. Hydrellia (Ephydridae) and some
other acalyptrate Diptera reared from Potamoge-
ton. Annals of the Entomological Society of
America 43: 374-398.
Busacca, J. D. and B. A. Foote. 1978. Biology and
immature stages of two species of Notiphila, with
notes on other shore flies occuring in cattail
marshes (Diptera: Ephydridae). Annals of the En-
tomological Society of America 71: 457—46.
Dacey, J. W. H. 1980. Internal winds in water lilies:
an adaptation for life in anaerobic sediments. Sci-
ence 210: 1017-1019.
Dacey, J. W. H. 1981. Pressurized ventilation in the
yellow water lily. Ecology 62: 1137-1147.
Deonier, D. L. 1972. Observations on mating, ovi-
position, and food habits of certain shore flies
(Diptera: Ephydridae). Ohio Journal of Science
72: 22-29.
Deonier, D. L., W. N. Mathis, and J. T. Regensburg.
1978. Natural history and life-cycle stages of No-
tiphila carinata (Diptera: Ephydridae). Proceed-
ings of the Biological Society of Washington 91:
798-814.
Eastin, W. C. and B. A. Foote. 1971. Biology and
immature stages of Dichaeta caudata (Diptera:
Ephydridae). Annals of the Entomological Society
of America 64: 271-279.
Grosse, W. and J. Mevi-Schutz. 1987. A beneficial gas
transport system in Nymphoides peltata. American
Journal of Botany 74: 947-952.
Hartley, J. C. 1958. The root-piercing spiracles of the
larva of Chrysogaster hirtella Loew (Diptera:
Syphridae). Proceedings of the Royal Entomolog-
ical Society of London 33: 81-87.
Hinton, H. E. 1953. Some adaptations of insects to
environments that are alternately dry and flooded,
with some notes on the habits of the Stratiomyi-
dae. Transactions of the Society for British Ento-
mology 11: 209-227.
Hinton, H. E. 1968. Respiratory systems of insect
eggs shells. Annual Review of Entomology 14:
343-368.
Hoglund, J. and R. V. Alatalo. 1995. Leks. Princeton
University Press, Princeton, N. J. xiii + 248 p.
Houlihan, D. EF 1969a. The structure and behavior of
Notiphila riparia and Erioptera squalida, two
root-piercing insects. Journal of Zoology 159:
1517-1536.
Houlihan, D. F 1969b. Respiratory physiology of the
larva of Donacia simplex, a root-piercing beetle.
Journal of Insect Physiology 15: 1517-1536.
Huryn, A. D. 1984. New Notiphila (Diptera: Ephyd-
ridae) from the Okefenokee Swamp, Georgia. Pro-
ceedings of the Entomological Society of Wash-
ington 86: 942-945.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Huryn, A. D. 1987. A new species of Notiphila (No-
tiphila) (Diptera: Ephydridae) from Ohio. Pro-
ceedings of the Entomological Society of Wash-
ington 89: 322-324.
Keilin, D. 1944. Respiratory systems and respiratory
adaptations in larvae and pupae of Diptera. Para-
sitology 36: 1—67.
Marshall, S. A. 1982. Techniques for collecting and
handling small Diptera. Proceedings of the Ento-
mological Society of Ontario 113: 73-74.
Mathis, W. N. 1979. Studies of Notiphilinae (Diptera:
Ephydridae), 1: Revision of the Nearctic species
of Notiphila Fallén, excluding the caudata group.
Smithsonian Contributions to Zoology 287: 1—
lila
Mathis, W. N. and T. Zatwarnicki. 1995. World Cat-
alog of Shore Flies (Diptera: Ephydridae). Assoc.
Publ., Gainesville, FL. 423 p.
Mevi-Schutz, J. and J. Grobe. 1988. A two-way gas
transport system in Nelumbo nucifera. Plant Cell
Environment 11: 27—34.
Mitter, C., B. D. Farrell, and B. Wiegmann. 1988. The
phylogenetic study of adaptive radiation: has phy-
tophagy promoted insect diversification? Ameri-
can Naturalist 132: 107-128.
Mitsch, W. J. and J. G. Gosselink. 1992. Wetlands.
Van Nostrand Reinhold, New York.
Miiller, G. W. 1922. Insektenlarven an Wurzeln von
Wasserplanzen. Mitteilungen des Naturwissen-
schaftlichen Vereines fur Neu-Vorpommern und
Rugen in Greifswald 48/49: 30-50.
Simpson, K. W. 1976. Shore flies and brine flies (Dip-
tera: Ephydridae), pp. 465—492. Jn Cheng, L. ed.,
Marine Insects. North Holland Publishing Com-
pany, Amsterdam.
Southwood, T. R. E. 1978. Ecological Methods. Chap-
man and Hall, New York. xxii + 424 p.
Todd, J. L. and B. A. Foote. 1987. Spatial and tem-
poral distribution of shore flies in a freshwater
marsh (Diptera: Ephydridae). Proceedings of the
Entomological Society of Washington 89: 448—
457.
Van der Velde, G. and C. M. Brock. 1980. The life
history and habits of Notiphila brunnipes Robi-
neau-Desvoidy (Diptera: Ephydridae), an auteco-
logical study on a fly associated with nymphaeid
vegetations. Tijdschrift voor Entomologie 123:
105-127.
Varley, G. C. 1937. Aquatic insect larvae that obtain
oxygen from the roots of plants. Proceedings of
the Royal Entomological Society of London 12:
55-60.
Varley, G. C. 1939. On the structure and function of
the hind spiracles of the beetle Donacia (Cole-
optera: Chrysomelidae). Proceedings of the Royal
Entomological Society of London 14: 115-123.
Whitman, R. L., J. M. Inglis, W. J. Clark, and R. W.
Clary. 1983. An inexpensive and simple elutria-
VOLUME 99, NUMBER 3 559
tion device for separation of invertebrates from Alpine, J. F, B. V. Peterson, G. E. Shewell, H.
sand and gravel. Freshwater Invertebrate Biology J. Teskey, J. R. Vockeroth, and D. M. Wood,
5: 159-163. coord. Manual of Nearctic Diptera. Research
Wirth, W. W., W. N. Mathis, and J. R. Vockeroth. Branch, Canada Department of Agriculture
1987. Ephydridae, pp. 1027-1047. In Mc- Monograph 28. Ottawa.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 560-573
DESCRIPTIONS OF FOUR NEW SPECIES OF CYNIPID GALL WASPS OF
THE GENUS NEUROTERUS HARTIG (HYMENOPTERA: CYNIPIDAE) WITH
REDESCRIPTIONS OF SOME KNOWN SPECIES FROM THE EASTERN
UNITED STATES
GEORGE MELIKA AND WARREN G. ABRAHAMSON
Department of Biology, Bucknell University, Lewisburg, PA 17837, U.S.A.; (GM) per-
manent address: Savaria Museum, Kisfaludy S.U.9, Szombathely, 9701 Hungary.
Abstract.—Four new species of cynipid gall wasps of the genus Neuroterus Hartig from
the eastern United States are described: N. archboldi Melika and Abrahamson, N. chap-
manii Melika and Abrahamson, N. christi Melika and Abrahamson, and N. weldi Melika
and Abrahamson. Four common eastern United States species of Neuroterus are rede-
scribed and four species are newly synonymized: N. quercusbatatus (Fitch) (Cynips nox-
iosus Bassett, new synonymy); N. quercusirregularis (Osten Sacken) (Cynips quercus-
majalis Bassett, new synonymy); N. quercusminutissimus (Ashmead); and N. quercus-
verrucarum (Osten Sacken) (Cynips flocossus Bassett and Neuroterus exiguissimus Bas-
sett, new synonymies).
Key Words:
ology
Neuroterus Hartig, 1840, is a holarctic
genus with numerous representatives in the
Old World (Eurasia) and North America.
Burks (1979) listed 52 species for America
north of Mexico, 33 of which are restricted
to the eastern United States. This genus is
easily distinguished from other genera of
oak gall-inducing cynipids by the absence
of a scutoscutellar suture; usually smooth
and thin body with a delicate coriaceous or
alutaceous sculpture on the thorax; radial
cell of fore wing long and narrow; and usu-
al absence of notauli. All known species of
this genus cause galls only on oaks of the
subgenus Lepidobalanus, except N. chry-
solepis Lyon which is associated with
Quercus chrysolepis Liebm. of the subge-
nus Protobalanus (Lyon 1984). The struc-
ture of Neuroterus galls is usually more
primitive than those of other genera, and
they lack highly specialized tissues and lay-
Cynipidae, gall wasps, Neuroterus, taxonomy, morphology, distribution, bi-
ers. Furthermore, the walls of the larval cell
usually are incorporated into the outer tis-
sues of the gall and do not separate from
the gall’s wall when the insect matures.
Numerous species of Neuroterus were
described from the eastern United States
before the end of the previous century, and
very often the only differences among the
described species were the host oaks from
which the galls were collected. Further-
more, the descriptions of the species are in-
complete and insufficient. Thus, eight com-
mon species of Neuroterus known from the
eastern United States are very difficult to
identify, either on the basis of the galls or
adults. These are N. quercusirregularis (Os-
ten Sacken) and N. quercusmajalis (Bassett);
N. quercusbatatus (Fitch) and N. noxiosus
(Bassett); and N. exiguissimus Bassett, N. flo-
cossus (Bassett), N. quercusminutissimus
(Ashmead), and N. gquercusverrucarum (Os-
VOLUME 99, NUMBER 3
ten Sacken). The descriptions and diagnostic
characters for separation of these species giv-
en by various authors (Ashmead 1885a,
1885b, 1887; Bassett 1864, 1881, 1900; Fitch
1859; Osten Sacken 1861, 1865) hardly allow
discrimination of all species. Examination of
types of these species indicates that some are
synonyms. Kinsey (1923), in his revision of
Neuroterus, treated N. exiguissimus, N. flo-
cossus, and N. minutissimus as varieties of
N. quercusverrucarum, N. noxiosus as a va-
riety of N. quercusbatatus, and N. quercus-
majalis as a variety of N. quercus-
irregularis. He was correct in splitting these
species, except N. quercusminutissimus
which, in our opinion, is a distinct species.
However, Kinsey’s synonymies did not fol-
low the International Code of Zoological
Nomenclature. Furthermore, his use of “‘va-
riety” and ‘‘form”’ makes his classification
confusing and difficult to use. Nevertheless,
we give credit to Kinsey for recognizing the
similarities among the above-listed species.
Thus, redescriptions, diagnostic characters,
taxonomic comments, and data on distri-
bution and biology for these eight species
are given.
We follow the current terminology for
morphological structures (Eady and Quin-
lan 1963, Fergusson 1995, Gibson 1985,
Menke 1993, Ritchie and Peters 1981, Ron-
quist and Nordlander 1989). The term “‘tho-
rax’’ used here includes the propodeum and
thus is equivalent to the “‘mesosoma”’ or
‘“mesosoma+metasoma”’ of the American
literature. Abbreviations for fore wing ve-
nation follow Ronquist and Nordlander
(1989). Measurements and abbreviations
used here include: Fl—F12, first and sub-
sequent flagellomeres; POL (post-ocellar
distance), the distance between the inner
margins of the posterior ocelli; and OOL
(odellar-ocular distance), the distance from
the outer edge of a posterior ocellus to the
inner margin of the compound eye.
Depositories for specimens are the Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C.
(USNM) and the American Museum of
561
Natural History, New York, New York
(AMNB).
Neuroterus archboldi Melika and
Abrahamson, NEW SPECIES
(Figs. 1-6)
Diagnosis.—No Neuroterus galls known
from the United States have galls similar to
N. archboldi (Figs. 5, 6).
Description.—Bisexual female: Dark
brown to black, with frons, clypeus and
mouthparts of some specimens yellow
brown. Head from above about twice as
wide as long, slightly broader than thorax,
gena not broadened behind eye (Fig. 1); in-
terocular space punctate, black, broader
than high; vertex finely coriaceous; distance
between antennal sockets shorter than dis-
tance between socket and inner margin of
eye; no distinct carina between antennal
sockets; frons punctate, broader than high,
yellow brown or yellow, with few scattered
pale setae; clypeus yellow, rounded, its
apex distinctly emarginate; anterior tento-
rial pits deep; malar space short, with faint
malar groove. Antenna 13-segmented, yel-
low, as long as head and thorax together,
with pale dense setae, basal 4 segments
lighter than rest; Fl slightly shorter than
pedicel and scape together, twice as long as
F2 (Fig. 2). Scutum rounded, only very
slightly broader than long, smooth and
shining, very finely coriaceous, without
trace of notauli, anterior parallel, and par-
apsidal lines; posterior margin slightly
emarginate on both sides of transverse
groove at base of scutellum; groove deep,
smooth. Scutellum slightly longer than
broad, smooth, shining, finely coriaceous
with very few scattered short, pale setae.
Pronotum, mesepisternum, and sides of
propodeum finely punctate, lighter than
scutum and scutellum. Medial part of pro-
podeum uniformly sculptured, without ca-
rinae. Fore wing 1.5 mm long, uniformly
and densely hairy, longer than body, with
cilia on margins, with light smoky spot on
M at junction of Cu,; veins thick, brown,
Rs+M reaches M;; areolet distinct, triangu-
562
larly rounded (Fig. 4). Legs, including cox-
ae pale yellow, semitranslucent, pretarsus
dark brown to black; claws without tooth.
Gaster darkish brown, smooth, in dry
shrunken specimens, nearly same height
and length; tip of ovipositor slightly curved,
ventral spine of hypopygium visible later-
ally, with short sparse pale setae. Length,
1.3-1.4 mm. Male: Color lighter than fe-
male, except for head. Eye slightly larger
than in female, interocular space black,
punctate, with duller sculpture than finely
coriaceous vertex. Anterior tentorial pits in-
distinct (unusual for males of species that
induce integral leaf galls; usually males
with very deep tentorial pits). Antenna
14-segmented, Fl longer than pedicel and
scape together, only very slightly curved
and extended in posterior part (Fig. 3). Fore
wing slightly darker than in female, larger,
length 2.0 mm. Petiole very distinct, long.
Length, 1.3 mm.
Types.—Holotype 2; allotype d6 from
Archbold Biological Station, Lake Placid,
Highlands Co., Florida, on Q. chapmanii
Sarg., 28 March 1995, emerged 30 March
1995. Also 25 @ and 4 ¢d paratypes. Ho-
lotype, allotype, 4 2 paratype, 1 d para-
type, and galls in the USNM; 5 @& para-
types, | ¢ paratype and galls in AMNH; 15
2 paratype, 2 ¢ paratypes, and galls in G.
Melika private collection.
Etymology.—Named in honor of Mr.
Richard Archbold, founder of the Archbold
Biological Station.
Distribution.—Florida (Archbold Biolog-
ical Station, Lake Wales Ridge, Highlands
Co.; Jonathan Dickinson State Park, Martin
Co.).
Biology.—Only the bisexual generation
is known, and the only known host for WN.
archboldi is Quercus chapmanii. This spe-
cies induces blister-like parenchyma thick-
enings which are visible on both sides of
the leaf but protrude more on the underside
of the leaf. The gall is rounded, monothal-
amous, |.5-2.0 mm in diameter, 1.0-1.5 mm
high, green or whitish green, very slightly
lighter than the leaf, and drier than the suc-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
culent polythalamous N. quercusirregularis
galls (Figs. 5, 6). The larval cell is situated
on the underside of the leaf, not separated
from the outer layers of the gall. There are
18 to 25 or more galls per leaf. After adults
emerge, the galls shrink and wrinkle and
dry out and drop. Consequently, rounded
holes remain in the leaf. We observed these
galls actively growing for the first time on
9 March 1995. Adults emerged the second
and third week of March into April. They
were very common leaf galls on Q. chap-
manii at the Archbold Biological Station.
Simultaneous with this species, N. quercus-
irregularis galls could develop on the same
leaf, but they are easily distinguishable
from those of the former on the basis of
galls. Our efforts to rear the alternate gen-
eration in cages containing emerged wasps
on Chapman oaks were unsuccessful.
Neuroterus chapmanii Melika and
Abrahamson, NEW SPECIES
(Figs. 7-11)
Diagnosis.—Three northeastern United
States species of Neuroterus are similar to
this new species on the basis of galls: WN.
dubius Bassett (egg-shaped capsule galls on
the edge of leaves on aments (Kinsey
1923)); N. exiguus Bassett (galls a fleshy
enlargement of the staminate axis (Weld
1959), or anther galls but very succulent
and shrivel up and disappear after adults
emerge (Bassett 1900)); and N. pallidus
Bassett (galls are in clusters, usually on the
end of catkins). Adults of all three men-
tioned species have character states that dif-
fer from N. chapmanii. Neuroterus dubius
has notauli and a coriaceous scutellum
while the other three species lack notauli
and have a smooth, shiny scutellum. Neu-
roterus exiguus has deep tentorial pits and
the head in front view is nearly as high as
broad, while in N. chapmanii and N. palli-
dus the head in front view is broader than
high with shallow tentorial pits. In N. chap-
manii, the head has a faint malar groove
and the POL:OOL is as 3.5:2.0, while the
VOLUME 99, NUMBER 3 563
Figs. 1-11. 1-6, Neuroterus archboldi. 1, Female head, front view. 2, Antenna of female. 3, Antenna of
male. 4, Fore wing of female. 5, Typical arrangement of galls on leaf (1X). 6, Saggital section of gall showing
larval chamber (10X). 7-11, N. chapmanii. 7, Female head, front view. 8, Antenna of female. 9, Antenna of
male. 10, Fore wing of female. 11, Shape of a single gall (30x).
564
head of N. pallidus lacks a malar groove
and the POL:OOL is as 3.0:2.0.
Description.—Bisexual female: Dark
brown to black. Head from above about
twice as wide as long, broader than thorax,
gena not broadened behind eye; interocular
space coriaceous, broader than high; dis-
tance between antennal sockets same as dis-
tance between antennal socket and eye;
frons coriaceous, broader than high, with
densely and uniformly distributed short pale
setae; clypeus rounded, its apex lighter,
emarginate; malar space very short, with
faint groove; mouthparts yellowish brown
(Fig. 7). Antenna 12- or 13-segmented
(some specimens with suture between 12th
and 13th segments indistinct), pale yellow,
with short pale dense setae; longer than
head and thorax together; Fl as long as
pedicel and scape together, two times lon-
ger than F2 (Fig. 8). Scutum rounded,
slightly longer than broad, smooth and
shiny, very finely coriaceous, without trace
of notauli, anterior parallel and parapsidal
lines. In specimens with shrunken and col-
lapsed thorax, median line and anterior par-
allel lines appear as dark lines. Posterior
margin of scutum slightly emarginate on
both sides from transverse groove at base
of scutellum; groove large, smooth, in-
curved. Scutellum smooth, shining, very
finely coriaceous with uniformly distributed
short pale setae; slightly elongated. Prono-
tum and mesepisternum finely coriaceous,
slightly lighter than scutum. Median part of
propodeum uniformly coriaceous, some
specimens with distinct median carina,
some without carina. Fore wing longer than
body, uniformly and densely pubescent,
with cilia on margins, without smoky spots;
veins thick, brown except paler Rs+M
which reaches M; areolet triangular, large;
length 1.5 mm (Fig. 10). Legs, including
coxae, pale, semitranslucent; basal part of
coxae and femora dark brown; pretarsus
dark brown or black; claws without tooth.
Gaster black, smooth, in dry specimens
shrunken and wrinkled, higher than long;
Ovipositor straight, tip not hooked; ventral
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
spine of hypopygium with short sparse pale
setae, prominent, visible laterally. Length,
1.1 mm. Male: Color lighter than female.
Eye considerably larger than in female, an-
terior tentorial pits deep; malar space short-
er than in female; antenna 14-segmented,
Fl longer than in female, very slightly
curved and extended posteriorly (Fig. 9). In
specimens with shrunken and collapsed tho-
rax, median line and anterior parallel lines
visible as dark lines. Fore wing longer than
in female, 1.8 mm. Petiole long. Length, 1.2
mm.
Types.—Holotype @, allotype d, 17 2
and 7 6 paratypes. Archbold Biological
Station, Lake Placid, Highlands Co., Flori-
da, 6 March 1995, emerged 8 March 1995.
Holotype, allotype, 4 2 and 2 ¢ paratypes
and galls in the USNM; 4 @ and 2 ¢ para-
types and galls in AMNH; 9 @ and 3 6
paratypes and galls in G. Melika private
collection.
Etymology.—Named from the host,
Quercus chapmanii.
Distribution.—Florida (Archbold Biolog-
ical Station, Lake Placid, Highlands Co.;
Jonathan Dickinson State Park, Martin
Co.).
Biology.—Only the bisexual generation
is known. Galls are on catkins of Quercus
chapmanii, are ovoid, thin walled, up to 1.5
mm long, and with a groove running from
one side to the other across the upper sur-
face thus resembling a closed purse (Fig.
11). The gall is covered with white pubes-
cence, especially on the top, and is mon-
othalamous. Galls are randomly scattered
along the staminate axis and perpendicular
to it. Sometimes two galls develop together.
The galls are surrounded by anthers. As the
gall and catkins mature, the color changes
from green to pale brown. They remain on
the catkins until emergence of the adult or
longer, dropping along with the catkins. In
Florida, on the Lake Wales Ridge, Chap-
man oak begins to flower the end of Feb-
ruary to early March. We found the gall for
the first time on 3 March 1995 when some
of the galls had emergence holes, likely
VOLUME 99, NUMBER 3
made by the gall-inducing wasps; also, pu-
pae and adult wasps were cut from the galls
on this date. The galls are found easily
throughout the period of Chapman oak
flowering. No inquilines were reared from
these galls, but some parasitoids belonging
to the Tetrastichinae (Hymenoptera: Eulo-
phidae) emerged. This is a very common
species on Q. chapmanii on the Lake Wales
Ridge. Our efforts to rear the alternate uni-
sexual generation were unsuccessful in
spite of using cages containing emerged
wasps on the branches of Chapman oaks.
Neuroterus christi Melika and
Abrahamson, NEW SPECIES
(Figs. 12-16)
Diagnosis.—The female appears most
similar to N. quercusirregularis but differs
by the parallel inner margins of the eyes,
shallow tentorial pits, smaller clypeus, and
ratio of the third antennal segment to the
first two together (1.7:1.0). In N. quercusir-
regularis, the eyes slightly converge inward
at the posterior part of the frons; the ante-
rior tentorial pits are deep, and the ratio of
the third antennal segment to the first two
segments combined is 1.3:1.0. The galls are
also quite similar in appearance to those of
N. quercusirregularis, but those of N. chris-
ti are strictly associated with Q. geminata
and Q. virginiana Mill., not nearly so suc-
culent, and usually of a regular cylindrical
shape and not so irregularly shaped as those
of N. quercusirregularis.
Description.—Bisexual female: Brown
to dark brown. Head from above about 3
times wider than long, broader than tho-
rax; gena not broadened behind eye, inner
margins of eyes parallel; interocular space
finely punctate, blackish brown to black,
broader than high; POL to OOL as 1.7:
1.0; distance between antennal sockets
smaller than between antennal socket and
inner margin of eye; antennal apodemes
distinctly depressed anteriorly; frons
lighter than vertex and body, coriaceous,
broader than high, with densely and uni-
formly distributed short pale setae, with
565
median elevation in form of broad carina
that widens posteriorly to width of clyp-
eus and reaches antennal sockets; clypeus
lighter than body and vertex, rounded,
very slightly emarginated posteriorly; ten-
torial pits shallow; malar space short, with
distinct malar groove (Fig. 12). Antenna
13-segmented, lighter than body, Fl
slightly longer than pedicel and scape to-
gether and 2 times longer than F2 (Fig.
13). Scutum rounded, slightly longer than
broad, smooth and shining, very finely co-
riaceous, without trace of notauli, anterior
parallel, median, and parapsidal lines;
however, possibly indicated by darker
lines; with scattered very short pale setae;
posteriorly emarginate on both sides from
transverse groove at base of scutellum.
Scutellum slightly longer than broad,
smooth, shiny, very finely coriaceous,
with very few scattered short pale setae;
transverse groove anteriorly very distinct,
deep, smooth, slightly incurved. Prono-
tum and mesepisternum shining, finely
coriaceous. Medial part of propodeum
finely punctate, without carinae, shiny and
smooth on both sides. Fore wing hyaline,
with cilia on margins, with smoky spot
along M distally from areolet and with a
very small darker smoky spot on junction
of M+Cu, to M. Fore wing 1.9-2.0 mm
long, slightly longer than body, areolet
present, closed; Rs+M lighter than other
veins (Fig. 15). Coxae and trochanters
pale yellow, semitranslucent, sometimes
tips of coxae brown; femora and tibia
brown, tarsi yellowish brown, pretarsi
blackish brown; claws without tooth. Gas-
ter dark brown to black, smooth, higher
than long; tip of ovipositor hooked; ven-
tral spine of hypopygium prominent, vis-
ible laterally, with pale scattered setae.
Length, 1.7 mm. Male: Color lighter than
female. Eye larger than in female; antenna
14-segmented; F1 longer than pedicel and
scape together, slightly curved and ex-
tended posteriorly (Fig. 14). Fore wing
2.2 mm long; body 1.6 mm long.
Types.—Holotype ¢, Bok Tower Gar-
566
dens, Polk Co., Florida, on Q. geminata,
26 March 1995, emerged 26-27 March
1995. Allotype 6, Lake Manatee State
Recreation Area, Manatee Co., Florida, on
Q. geminata, 9 April 1995, emerged 10
April 1995. 10 2 and 5 ¢ paratypes. Ho-
lotype, allotype, 3 2 and 2 d paratypes
and galls in the USNM; 3 2 and 1 ¢ para-
types and galls in AMNH; 4 2 and 2 ¢
paratypes and galls in G. Melika private
collection.
Other material examined.—13 2 and 9
6, in addition to the two above-mentioned
localities, from Buck Island Ranch, 13 km
SW Brightly (Archbold Biological Station
property), Highlands Co., Florida, on Q.
virginiana, coll. 17 March 1995, emerged
26 March 1995.
Etymology.—Named in honor of Ms.
Christy Raye Abrahamson, who provided
continuous support during our seven
months of research based at the Archbold
Biological Station, and who first found this
species.
Distribution.—Florida (Archbold Biolog-
ical Station and 13 km SW Brighton, Buck
Island Ranch, Highlands Co.; Lake Mana-
tee Recreation Area, Manatee Co.; Bok
Tower, Polk Co.; Eglin Air Force Base,
Okaloosa and Walton cos.; Wakull Spring
State Park, Wakulla Co.).
Biology.—Only the bisexual generation
is known. The host oaks are Quercus gem-
inata and Q. virginiana. The galls are leaf
parenchyma thickenings, equally protrud-
ing on both sides of the leaf along one side
of the main vein, green when young but
turning brownish green when mature, po-
lythalamous, generally with four to six lar-
val cells per gall, about 12.0-15.0 mm
long, 3.0-4.5 mm wide, and 3.0-4.5 mm
high, and usually of a regular cylindrical
form. The walls of the inner cells are whit-
ish green and the surface of the gall is
smooth, without hairs (Fig. 16). There are
usually one or two galls per leaf. Adults
usually emerge from the upper side. The
first galls appear in mid-March, and the
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
adults emerge the last week of May and
beginning of June.
Neuroterus weldi Melika and
Abrahamson, NEW SPECIES
(Figs. 17-21)
Diagnosis.—This species is related to N.
quercusbatatus (Fitch), but the gena are
broadened behind the eyes; a distinct carina
on the vertex reaches between the antennal
sockets; the area around the antennal sock-
ets is depressed; the medial carina on the
frons is distinct only in the lower half; the
ratio between the first four antennal seg-
ments (1+-2:3:4) is 1.16:1.03:0.63) andoan
dried specimens, the height and length of
the gaster is similar, but in some specimens
the gaster is longer than high. In N. quer-
cusbatatus, the genae are not broadened be-
hind the eyes; the carina on the vertex is
less distinct; the area around the antennal
sockets is less depressed; the median carina
on the frons is longer and reaches the an-
tennal sockets; the ratio between the first
four antennal segments (1+2:3:4) is 0.93:
0.77:0.57; and, in dried specimens, the gas-
ter is higher than long. The phenology and
gall structure of both species also differ (see
biology section of both species).
Description.—Unisexual female: Brown
to red brown, with dark brown to black gas-
ter. Head piceous, from above about 2 times
wider than long, broader than thorax, gena
broadened behind eye; interocular space co-
riaceous, broader than high; distance be-
tween antennal sockets smaller than be-
tween eye and antennal socket; distinct ca-
rina on vertex reaching antennal sockets;
frons coriaceous, broader than high, with
densely and uniformly distributed short pale
setae; median carina on frons indistinct,
never extending between antennal sockets
(like that in N. quercusbatatus); malar
space well developed with deep malar
groove; frons and clypeus lighter than rest
of head, piceous; mouthparts of same color
as frons and clypeus; tip of mandible black
(Fig. 17). Antenna 13-segmented, longer
than head and thorax together, pedicel and
VOLUME 99, NUMBER 3 567
Figs. 12-21. 12-16, Neuroterus christi. 12, Female head, front view. 13, Antenna of female. 14, Antenna
of male. 15, Fore wing of female. 16, Shape and usual location of gall (3.5X). 17-21, N. weldi. 17, Female
head, front view. 18, Antenna of female. 19, Fore wing of female. 20, Ventral spine of hypopygium of female,
ventral view. 21, Shape of gall (2.5).
568
scape flattened; scape slightly longer than
broad; first 3 segments piceous, rest dark
brown to black; ratio between first 4 seg-
ments .(1+-2:3:4)) as 1.116:1/03:0:63 rie:
18). Scutum rounded, slightly longer than
broad; brown, smooth and shining, finely
coriaceous, without trace of notauli, ante-
rior parallel, median, and parapsidal lines;
posterior margin slightly emarginate on
both sides from transverse groove at base
of scutellum; groove distinct, with shiny
bottom. Scutellum piceous, much lighter
than scutum, rounded, slightly longer than
broad, smooth, shiny, finely coriaceous,
with few short and uniformly distributed
pale setae. Pronotum and mesepisternum
finely coriaceous. Median part of propo-
deum uniformly finely coriaceously punc-
tate; sides similar. Fore wing translucent,
with uniform very short pubescence, with
cilia on margins, some specimens with only
trace of smoky spot at junction of M to Cu,
and on 2r-m; veins thick, brown, areolet
large, triangular, Rs+M reaches M (Fig.
19); length 1.9-2.1 mm. Legs uniformly red
brown, with black pretarsi; claws simple,
without tooth. Gaster black, smooth, slight-
ly longer than high; ovipositor tip hooked;
hypopygium with few short sparse setae
(Fig. 20). Length, 1.6-1.8 mm.
Comments.—Weld (1959) wrote of a
stem swelling found on Q. chapmanii, but
he never reared the adults and consequently
did not describe the species. The galls in-
duced by N. weldi are probably those Weld
described.
Types.—Holotype 2 and 17 2 paratypes.
Air Force Range, Avon Park, Highlands
Co., Florida, 9 February 1995, emerged 16
February 1995. Holotype and 4 paratypes
in the USNM; 4 paratypes in AMNH; 9
paratypes in the private collection of G.
Melika.
Etymology.—Named after the American
cynipidologist L. H. Weld who probably
first found the galls of this species.
Distribution.—Central and coastal sand-
ridges of south central Florida (Archbold
Biological Station, Lake Placid, Highlands
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Co.; Avon Park Air Force Range, High-
lands Co.; Jonathan Dickinson State Park,
Martin Co.).
Biology.—Only the unisexual generation
is known. The gall is cylindrical, mostly a
terminal stem swelling, covered with nor-
mal bark, on second year or older twigs and
branches only of Quercus chapmanii. It is
25.0-40.0 mm long, 6.0-9,0 mm in diame-
ter, and polythalamous (Fig. 21). Galls are
not woody, rather they are easily cut like
cheese. The leaf petioles are never involved
in gall formation. Galls persist on stems for
several years. Fully developed adults over-
winter in galls and emerge from the second
half of February into March of the follow-
ing spring. The first galls observed are
formed by the end of May.
Neuroterus quercusbatatus (Fitch)
Cynips Quercus-batatus Fitch 1859: 810.
Females and males (type examined).
Neuroterus batata Basset: Ashmead 1885a:
296; 303.
Neuroterus batatus form bisexualis Kinsey
1920: 334.
Neuroterus (Dolichostrophus) batatus vat.
batatus form bisexualis Kinsey 1923: 4.
Neuroterus quercusbatatus: Burks 1979:
1074.
Cynips noxiosus Bassett 1881: 108. Fe-
males, males, galls of both generations
(types examined). New synonymy.
Neuroterus noxiosus form vernalis Kinsey
1920: 337. Bisexual generation.
Diagnosis.—This species closely resem-
bles N. weldi; see diagnosis for that species.
Redescription.—Female: Brown to red
brown. Head from above 2 times broader
than long, broader than thorax, gena only
very slightly broadened behind eye; inter-
ocular space coriaceous, broader than high;
distance between antennal sockets nearly
same as distance between eye and antennal
socket; frons coriaceous, broader than high,
with uniformly distributed short pale setae;
median carina on frons reaches between an-
tennal sockets; malar space well developed
VOLUME 99, NUMBER 3
with deep groove; frons, clypeus, and
mouthparts of same color as rest of head,
in some specimens lighter; tip of mandible
dark brown to black. Antenna 13-segment-
ed, brownish yellow, sometimes pale yel-
low, slightly longer than head and thorax
together; 3 apical flagellomeres dark brown;
pedicel and scape flattened; scape slightly
longer than broad; ratio between first 4 seg-
ments (1+2:3:4) as 0.93:0.77:0.57. Scutum
rounded, slightly longer than broad, brown
black, smooth and shiny, finely coriaceous,
without trace of notauli, anterior parallel,
median, and parapsidal lines. Scutellum
dark brown to black, rounded, only very
slightly longer than broad, smooth, shiny,
finely coriaceous, with densely and uni-
formly distributed pale setae; posterior mar-
gin slightly emarginate on both sides from
transverse groove which is shiny. Prono-
tum, mesepisternum finely coriaceous. Me-
dial part of propodeum uniformly coria-
ceously punctate; sides similar. Fore wing
translucent, with uniform, very short pu-
bescence, with cilia on margins; veins
thick, brown, areolet large, triangular. Legs
brown; coxae, centers of femora, and hind
tibia somewhat darker; claws without tooth.
Gaster larger than thorax, brown, some-
times brown black, usually same color as
thorax and head, higher than long; ovipos-
itor tip hooked, ventral spine of hypopy-
gium with few short sparse white setae.
Length, 1.2-2.2 mm. Females of both gen-
erations quite similar, impossible to distin-
guish them on basis of morphological char-
acters. Male: Similar in color to female;
thorax red brown laterally; legs and antenna
uniformly yellow; gaster with petiole pice-
ous or lighter in color; eyes only slightly
enlarged; F1 not longer than that of female.
Distribution.—Ontario, Rhode Island,
Connecticut west to Illinois, Colorado,
south to Florida.
Biology.—Alternate bisexual and unisex-
ual generations are known. Both genera-
tions induce stem-swelling galls on Quer-
cus alba L. (Ashmead 1885a, Burks 1979).
In Florida, the galls of the bisexual gener-
569
ation also develop on Q. chapmanii and Q.
margaretta. The gall is a woody, elongate
stem swelling. It is polythalamous, with an
irregular shape and a surface that is covered
by normal bark, twisted, but in large part
cylindrical, tapering gradually to the stem
at both ends and up to 20 mm long and 8
mm wide in the bisexual form and 60 mm
long by 20 mm wide in the unisexual form.
Sometimes several galls can more or less
fuse, drying brown in bisexual forms, with
a whitish or purplish bloom in unisexual
forms. Internally hard and woody, the tissue
is little modified except by the larval cells
which are densely packed, each with a dis-
tinct but wholly inseparable lining. The uni-
sexual form develops on young stems, in-
volving petioles and leaf midveins; the bi-
sexual form is on older stems involving the
bases of the petioles. The unisexual form
begins to develop in mid-summer, forming
woody stem galls. The adults overwinter in
the galls and emerge in spring after the oaks
are actively growing. The galls of the bi-
sexual generation are less woody, usually
on younger twigs, and develop in late
spring to early summer. The adults emerge
in June and July.
Neuroterus quercusirregularis
(Osten Sacken)
Cynips q. irregularis Osten Sacken 1861:
65. Species described from one damaged
male (sex not certain) and one gall. Type
lost.
Neuroterus irregularis: Ashmead 1885a:
296, 304.
Neuroterus quercusirregularis: Burks 1979:
1074.
Cynips quercus-majalis Bassett 1864: 683.
Females, males, galls (types examined).
New synonymy.
Neuroterus majalis: Mayr 1881: 37.
Neuroterus (Dolichostrophus) irregularis
var. majalis: Kinsey 1923: 100.
Diagnosis.—This species is very closely
related to N. christi; see diagnosis of that
species. Neuroterus quercusirregularis is
570
associated with the Quercus chapmanii- Q.
margaretta-Q. stellata Wangenh. group,
while the closely related N. christi is found
only on Q. virginiana and Q. geminata.
Redescription.—Female: Head largely
dark brown to black, finely coriaceous;
gena not broadened behind eye; frons with
uniformly distributed short pale setae; eyes
enlarged, malar space very short with malar
groove; clypeus rounded, emarginated pos-
teriorly, slightly lighter than frons. Antenna
brown, yellow to whitish basally, 13-seg-
mented, Fl 2 times or more longer than F2.
Thorax black, only slightly longer than high
or wide. Scutum and scutellum smooth,
shiny, very finely coriaceous, without setae.
Pronotum and mesepisternum finely coria-
ceous. Fore wing densely pubescent, much
longer than the body, with cilia on margins;
areolet usually moderate, but variable in
size; with very light smoky spot a midpoint
of M. Legs entirely pale yellow or whitish,
semitranslucent; pretarsi black; claws with-
out tooth. Gaster black or very dark brown,
shining and smooth, higher than long.
Length, 1.5-2.0 mm. Male: Head dark
brown to black, lower part of frons lighter
than in female; eye considerably enlarged,
malar space very short. Antenna generally
lighter than in female, pale yellow or yel-
low white, with Fl much lengthened, dis-
tinctly curved, 2 times longer than both
pedicel and scape together and nearly 4
times longer than F2. Thorax considerably
longer than in female, light brown, laterally
yellow to whitish. Fore wing much longer
than body. Legs whitish, semitranslucent.
Male slightly larger than female.
Comments.—We found no evidence to
differentiate N. quercusirregularis from N.
quercusmajalis. Beutenmueller (1910)
wrote about N. quercusmajalis: ‘‘Allied to
N. q. irregularis in color ..., and the only
perceptible difference that I can find be-
tween the two species is in size.’’ Bassett
(1864) mentioned that is it quite similar to
Osten Sacken’s C. q. irregularis, but related
to other species of oak. Kinsey (1923) treat-
ed N. quercusmajalis as a variety of N. ir-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
regularis, and described coloration as the
only difference. Unfortunately, the type
male of N. quercusirregularis, originally
designated by Osten Sacken, is lost. Except
for slight difference in coloration, we found
no morphological differences between N.
quercusmajalis and N. quercusirregularis
in either sex. Consequently, we regard these
species as synonymous.
Distribution.—Ontario, New York, Mas-
sachusetts, Connecticut, New Jersey, Mary-
land, Virginia, west to Illinois, Missouri,
Oklahoma, Texas, and southward (Beuten-
mueller 1910). We found this species in
North Carolina (vicinities of Arapahoe,
Pamlico Co., on Q. stellata), and in Florida
(common, locally abundant, occurs every-
where the host plants grow: Jonathan Dick-
inson State Park, Martin Co.; Archbold Bi-
ological Station, Highlands Co.; Winegar-
ner’s property in the vicinity of De Funiak
Springs, Walton Co., and throughout the
panhandle and south central Florida).
Biology.—Only the bisexual generation
is known to induce integral leaf galls on
Quercus alba and Q. montana Willd. (Ash-
mead 1885a, Beutenmueller 1910, Kinsey
1923, Weld 1959), and Q. chapmanii and
Q. stellata (Burks 1979). We frequently
found this species on Q. margaretta as well.
According to Kinsey (1923), N. quercusir-
regularis var. albipleurae Kinsey induces
galls on Q. breviloba (in Texas only). The
gall is a leaf parenchyma thickening, with
the larval cells mostly deeply embedded,
and is polythalamous. It is usually elongate,
oval, as thick as wide, up to 5.0 mm wide
by 15.0 mm long, several galls often fusing,
green, very succulent, shriveling consider-
ably upon drying, and the larval cell has a
distinct but inseparable layer. The galls are
smooth and large when on Q. alba and the
pubescence is denser and the galls are
smaller when on Q. stellata, Q. margaretta,
or Q. chapmanii. Galls usually appear on
very young, unfolding leaves, very quickly
mature, and develop from April until early
June, earlier farther south. Adults emerge
from May through July.
VOLUME 99, NUMBER 3
Neuroterus quercusminutissimus
(Ashmead)
Cynips q. minutissima Ashmead 1885a: 7.
Females only.
Neuroterus minutissimus: Ashmead 1885b:
296.
Neuroterus (Diplobius) verrucarum vat.
minutissimus: Kinsey 1923: 72.
Diagnosis.—The sculpture of the prono-
tum in N. quercusverrucarum is duller than
in N. quercusminutissimus. The gaster in N.
quercusminutissimus is sharply triangular,
considerably higher than long, and smaller
than the head and thorax together, while in
N. quercusverrucarum, the gaster is more
prolonged and nearly the same length as the
head and thorax together. The gall resem-
bles that of N. quercusverrucarum;, how-
ever, the cynipid species composition on Q.
virginiana-Q. geminata-Q. minima is very
specific. No other host plants have been re-
corded for the 14 species of oak cynipids
that occur on the closely related group of
oak species. Thus, knowledge of the host
species is very helpful in the identification
of species. Neuroterus quercusverrucarum
and N. quercusminutissimus are both very
common in Florida; however, the former
species is associated with Q. chapmanii, Q.
stellata, and Q. margaretta, while the latter
species is hosted by Q. virginiana, Q. gem-
inata, and Q. minima.
Redescription.—Female: Entire body
black, only clypeus and mandible light
brown. Head from above 2 times wider than
long, broader than thorax, gena only very
slightly broadened behind eye; interocular
space, vertex, and frons of same coriaceous
sculpturing; frons with uniformly densely
distributed short pale setae; malar space
moderately large, with faint groove. Anten-
na 13-segmented, yellowish brown, F1 lon-
gest flagellomere, but only slightly longer
than F2. Scutum and scutellum both round-
ed, shiny, very finely coriaceous, with very
few short pale setae on scutum and with
uniformly distributed dense short white se-
tae on scutellum; without trace of notauli,
3) 7/3!
anterior parallel, median, and parapsidal
lines; transverse groove at base of scutellum
large, with smooth, shiny bottom. Pronotum
and mesepisternum same color as scutum,
very finely coriaceous. Fore wing translu-
cent, with cilia on margins, longer than
body; veins thick, yellowish, only Rs+M
much paler; areolet triangular, distinct. Legs
yellowish brown; coxae, femora, and tibiae
infuscated along upper surface; claws with-
out tooth. Gaster black, shiny, higher than
long, sharply triangular in outline; ovipos-
itor straight, tip not hooked; ventral spine
of hypopygium prominent, visible laterally.
Length, 0.5-1.3 mm.
Distribution.—Florida. Common every-
where Quercus geminata, Q. minima, and/
or Q. virginiana grows.
Biology.—Only the unisexual generation
is known. This species induces small, 1.25-
2.50 mm, detachable galls, covered with
coarse, light brown mossy pubescence, on
the underside of leaves in numbers in the
fall on Quercus geminata, Q. virginiana,
and Q. minima. In Florida, adults emerge
early in the spring, but the galls do not de-
velop until mid- to late summer. The fully
grown adults overwinter in the galls.
Neuroterus quercusverrucarum
(Osten Sacken)
Cynips quercus verrucarum Osten Sacken
1861: 62. Females and galls (types ex-
amined).
Cynips verrucarum: Osten Sacken 1865:
354.
Neuroterus verrucarum: Ashmead 1885a:
296, 304.
Neuroterus quercusverrucarum: Burks
1979: 1075.
Neuroterus exiguissimus Bassett 1900: 332.
Females and galls (types examined). New
synonymy.
Cynips flocossa Bassett 1881: 111. Females
and galls (types examined). New synon-
ymy.
Neuroterus verrucarum var. pernotus Kin-
sey 1923: 74.
Neuroterus flocossus: Burks 1979: 1073.
S72
Diagnosis.—See Neuroterus quercusmi-
nutissimus.
Description.—Female: See description
and diagnosis for N. quercusminutissimus.
Length, 0.7-1.5 mm.
Comments.—We found no morphologi-
cal differences in adults, galls, and phenol-
ogy of N. quercusverrucarum, N. exiguis-
simus, and N. flocossus; consequently we
treat them as synonyms. Kinsey (1923) de-
scribed ten varieties of this species, one of
which, N. verrucarum var. restrictus Kin-
sey, was from Florida on Quercus chap-
manii. He indicated that this variety was
collected also on Q. margaretta and Q.
geminata. In our opinion, the latter refer-
ence concerns N. quercusminutissimus.
Distribution.—Ontario, New York, New
Jersey, Connecticut, west to Illinois, Okla-
homa, Texas, south to Florida.
Biology.—Only the unisexual generation
is known. This species induces detachable
leaf galls which are small, smooth, seed-
like larval cells, hard, thin-walled, and cov-
ered with a dense flattened yellowish brown
mass of wool that dries brown. The gall is
monothalamous, attached to the underside
of the leaf by a small point to the leaf vein,
separable, but the leaf is depressed at the
point of attachment. The average diameter
of a gall is 2.0 mm by 1.0 mm high. The
galls are usually numerous, covering the
whole underside of the leaf and are found
on Quercus alba, Q. bicolor Willd., Q.
chapmanii, Q. margaretta, and Q. stellata.
Viereck (1916) also recorded Q. macrocar-
pa Michx. as a host, but this must be con-
firmed. Galls begin to develop in September
(Weld 1959). According to Kinsey (1923),
the galls appear after mid-summer in Au-
gust and the larvae do not mature until late
in the fall. Adults emerge the next spring in
February to March.
ACKNOWLEDGMENTS
We express our deepest appreciation to J.
Fitzpatrick for his support of our work at
the Archbold Biological Station, M. Deyrup
for providing laboratory space and support,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and C. Abrahamson for extensive field and
technical assistance. We also thank R. Lyon
for suggestions and comments. We thank J.
Abrahamson, R. Bowman, G. Cs6ka, R.
Hammer, A. Johnson, I. Kralick, R. Peet, R.
Roberts, P. Schmaltzer, A. Schotz, C. Wine-
garner, and M. Winegarner for field and
technical assistance. Our special thanks to
A. S. Menke for his valuable suggestions
and support during our work at the National
Museum of Natural History, Smithsonian
Institution, Washington, DC. Support was
provided to GM and WGA by Bucknell
University’s David Burpee endowment, the
Archbold Biological Station, a National
Museum of Natural History, Smithsonian
Institution grant to GM, and NSF grant
BSR-9107150 to WGA.
LITERATURE CITED
Ashmead, W. H. 1885a. On the cynipidous galls of
Florida with descriptions of new species. Trans-
actions of the American Entomological Society
12: 5-9.
Ashmead, W. H. 1885b. A bibliographical and syno-
nymical catalogue of the North American Cyni-
pidae, with description of new species. Transac-
tions of the American Entomological Society 12:
291-304.
Ashmead, W. H. 1887. On the cynipidous galls of
Florida, with descriptions of new species and syn-
opses of the described species of North America.
Transactions of the American Entomological So-
ciety 14: 125-158.
Ashmead, W. H. 1903. Classification of the gall-
wasps and the parasitic cynipoids, or the super-
family Cynipoidea. III. Psyche 10: 140-155.
Bassett, H. E 1864. Descriptions of several new spe-
cies of Cynips, and a new species of Diastrophus.
Proceedings of the Entomological Society of Phil-
adelphia 3: 679-691.
Bassett, H. E 1881. New Cynipidae. Canadian Ento-
mologist 13: 92-113.
Bassett, H. EK 1900. New species of north American
Cynipidae. Transactions of the American Ento-
mological Society 26: 310-336.
Beutenmueller, W. 1910. The North American species
of Neuroterus and their galls. Bulletin of the
American Museum of Natural History 28: 117—
136.
Burks, B. D. 1979. Superfamily Cynipoidea, pp.
1045-1107. In Krombein, K. V., P. D. Hurd, Jr.,
D. R. Smith, and B. D. Burks, eds. Catalog of
Hymenoptera in America North of Mexico. Vol-
VOLUME 99, NUMBER 3
ume 1. Symphyta and Apocrita. Smithsonian In-
stitution Press, Washington, DC.
Eady, R. D. and J. Quinlan. 1963. Handbooks for the
identification of British insects. London. VIII (Ia).
81 pp.
Fergusson, N. D. M. 1995. The cynipoid families, pp.
247-265. In Hanson, P. E., and I. D. Gauld eds.
The Hymenoptera of Costa Rica. Oxford, New
York, Tokyo, Oxford University Press.
Fitch, A. 1859 (1858). Fifth report on the noxious and
other insects of the state of New York. Transac-
tions of the New York State Agricultural Society
18: 781-854.
Gibson, G. A. P. 1985. Some pro- and mesothoracic
structures important for phylogenetic analysis of
Hymenoptera, with a review of terms used for the
structures. Canadian Entomologist 117: 1395-—
1443.
Kinsey, A. C. 1920. New species and synonymy of
American Cynipidae. Bulletin of the American
Museum of Natural History 42: 293-317.
Kinsey, A. C. 1923. The gall wasp genus Neuroterus
(Hymenoptera). Indiana University Studies 10: 1—
150.
Lyon, R. 1984. New Cynipid Wasps from California.
The Pan-Pacific Entomologist 60: 289-290.
Mayr, G. 1881. Die Genera der gallenbewohnenden
1/3)
Cynipiden. Jahresbericht der Rossauer Communal
Oberealschule, Wien 20: 1-38.
Menke, A. 1993. Notauli and parapsidal lines: just
what are they? Sphecos 24: 9-12.
Osten Sacken, C. R., von 1861. On the Cynipidae of
the North American oaks and their galls. Proceed-
ings of the Entomological Society of Philadelphia
1: 47-72.
Osten Sacken, C. R., von 1865. Contributions to the
natural history of the Cynipidae of the United
States and of their galls. Proceedings of the En-
tomological Society of Philadelphia 4: 331—380.
Ritchie A. J., and T. M. Peters. 1981. The external
morphology of Diplolepis rosae (Hymenoptera:
Cynipidae, Cynipinae). Annals of the Entomolog-
ical Society of America 74: 191-199.
Ronquist, E, and G. Nordlander. 1989. Skeletal mor-
phology of an archaic cynipoid, /balia rufipes
(Hymenoptera: Ibaliidae). Entomologica Scandi-
navica, Supplement 33: 1—60.
Viereck, H. L. 1916. Cynipoidea. In: Guide to the
insects of Connecticut. Part III. The Hymenoptera,
or wasp-like insects of Connecticut. Bulletin of
the State Geological and Natural History Survey
22: 361-443.
Weld, L. H. 1959. Cynipid galls of the eastern United
States. Ann Arbor, MI, Privately printed. 124 pp.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 574-582
OBSERVATIONS ON SOME HEMIPTERA/HETEROPTERA OF MACAU,
SOUTHEAST ASIA
EMMET R. EASTON AND WING-WAH PUN
(ERE) University of Macau, PO. Box 3001, Macau via Hong Kong; (WWP) Depar-
tamento de Servigos Agrarios de Camara Municipal das Ilhas, Coloane, Macau.
Abstract.—The insect species of Macau are believed to be quite diverse in numbers of
species due to trees and foliage representing the subtropical evergreen broadleaf forest as
well as remnants of the monsoon tropical rain forest. The absence of land under agricul-
tural production may account for the low numbers of some groups that are more common
in the temperate regions of the world such as the Anthocoridae, Miridae, or Lygaeidae.
Records of 59 species of true bugs are listed for the Portuguese territory of Macau of
which 32 are considered new records.
Key Words:
The true bugs locally have been little
studied since the time of J. C. W. Kershaw
who lived in Macau for seven years and
published a series of papers on local he-
miptera and homopteran fauna (Kershaw
and Kirkaldy 1908a, b, 1909a, b, c). The
territory of Macau consists of a peninsula
of land connected to the Guangdong Prov-
ince of mainland China as well as two is-
lands consisting of Taipa and Coloane
which are connected to the peninsula ei-
ther by bridges (Taipa, 2) or a causeway
(Coloane). It is most likely that Kershaw
carried out his observations on Macau in-
sects on the peninsular area as road access
to the islands was not available until
1974. Since that time urbanization has
eliminated most of the agricultural habi-
tats on the land area connected to main-
land China and the majority of our records
concern fauna of the islands where the ur-
banization process has been less intensive.
Notes are provided in this paper on certain
species in families the authors have ob-
served in Macau over the course of years
from 1989 to the present.
Macau, Hemiptera, Heteroptera, faunal list
MATERIALS AND METHODS
Insects were sampled during all months
of the year from an artificial light source
built in 1994 at the entrance of the inner
harbour on the Macau peninsula. The
presence of 18 high intensity lamps (1800
watt) illuminating the 40 metre tall granite
Friendship monument accounted for the
fair number of records of which Nezara
viridula (L.) (Pentatomidae) was found to
be the most abundant species present.
Other species were collected from the
walls of buildings (Taipa island) that are
normally illuminated at night while those
species not sensitive to hight were sam-
pled from trees and shrubs upon visual ex-
amination. Names of the plant hosts fol-
low the Macau Catalogue of Plants (Ca-
mara Municipal das Ilhas, Macau and In-
stituto de Investigacgao Cientifica Tropical,
Lisboa 1991). Representatives of the spe-
cies of Hemiptera named here are housed
in the entomology museum of the Agrar-
ian Services on Coloane island under the
curatorship of the second author.
VOLUME 99, NUMBER 3
LIsT OF SPECIES
HEMIPTERA
Acanthosomatidae
Dichobothrium nubilum (Dallas). 12
April 1994, Ip Tai leg, NEW RECORD,
Coloane Island. In mainland China it is
found in Hainan, Guangdong, Guangsi,
Hunan, Sichuan Provinces as well as Tibet;
also India, Taiwan and Japan.
Elasmostethus nobilus (Dallas). 18 April
1996, ERE leg, NEW RECORD, Friend-
ship monument on Macau peninsula Also
found-in Hong Kong according to Tai Lung
farm records.
Belostomatidae
Diplonychus rusticum FE, small water
buss 18..23, 25,28, 30:Mays, 1.42 June,.25
Aug. 1996, ERE leg, Friendship monument,
Macau peninsula; male with eggs, Leu-
ng-va leg, Coloane island, NEW RECORD.
Found in pond on reclaimed land. In Hong
Kong Dudgeon and Corlett (1994) report it
common in slow moving streams and
marshes. It is found in India, Burma, Sri
Lanka, Malaya, Sumatra, Java and Thailand
as well as Australia.
Lethocerus indicus (Lepeletier and Ser-
ville), giant water bug. 24 Jan.; 10 June
1994; 5 May; 10, 19 June 1995; 19 June
1996, ERE leg, NEW RECORD, Macau
peninsula. Hoffman (1933) studied its life
history in Guangzhou; eggs were laid on
upright vegetation in ponds. At lengths
ranging from 60-80 mm it is the largest
hemipteran known. Widespread in China it
is also found in Burma, Sn Lanka, India,
Java, Sumatra, Malaya and the Philippines.
Coreidae
Acanthocoris scaber (L.), brown coreid
bug. 11 July 1994, Cheong Chi Keong leg,
NEW RECORD, Coloane Island, ex Cap-
sicum frutescens, Ipomeae batatas, Lyco-
persicum esculatum and eggplant, Solanum
melongena. Hoffman (1931a) reported it
damaging eggplant, squash and peppers in
575
the Guangzhou area of SE China. Reported
also in India.
Cletus trigonus (Thunberg), slender rice
bug. 19 Sept. 1993. PWW leg, NEW REC-
ORD, Coloane Island, ex Amaranthus tri-
color, Chinese spinach. In addition to
Guangdong Province of China it also oc-
curs in India, Sr Lanka, Borneo and Phil-
ippines.
Gralliclava horrens (Dohrn), Oriental
pod bug. 17 Aug. 1993, Ip Tai leg, NEW
RECORD, Coloane Island, ex Arachis hy-
pogaea, groundnut. Dolling (1978) revised
the group and reported it from Hong Kong,
India, China (Fukien, Hainan), Burma,
Thailand, Borneo, Sulawezi, Philippines
and Taiwan.
Homeocerus unipunctatus Dallas. Date,
collector unknown. NEW RECORD, Co-
loane island, ex. Glycine max, Phaseolus
radiatus, vetch. Wu (1935) reported it also
in India, Sri Lanka, Burma, Malaya and
Java.
Leptocorisa acuta (Thunberg), coreid
rice bug. 12, 19 Nov. 1994, ERE leg,
Friendship monument, Macau peninsula. It
was reported earlier from Taipa island
(Easton 1992). Sands (1977) recorded it
from Southern Japan, Taiwan, Hong Kong,
Vietnam, India, Pakistan, Thailand, Indo-
nesia to Papua New Guinea, Samoa, Fiji
and Australia.
Mictis tenebrosa F, cassia bug. 9 Aug.
1992, Ng Wai Man leg, NEW RECORD,
Coloane Island, ex Arachis hypogoea and
Smilax china, greenbrier. In Hong Kong
there are records from Cassia fistula (Lee
and Winney 1981). O’Shea and Schaefer
(1980) reported it from India, Burma, Ma-
laya and Indonesia as well as China.
Notobitus meleagris (F.), bamboo coreid
or leaf-footed bug. 11 Oct. 1992, Ng Wai
Man leg, Coloane island; 2 nymphs, 12
adults, 30 Aug 1995, ERE leg, ex Dendro-
calmus pulverulentus bamboo in Lou Lim
Iok gardens, Macau peninsula. Reported
also in Hainan, China as well as India, Ma-
laya and Hong Kong.
Paradasynus spinosus Hsiao, green cor-
576
eid bug. 10 Nov. 1992, PWW leg, NEW
RECORD, Coloane Island. In Hong Kong
it has been found on Citrus sinensis and
Melia azedarach (Lee and Winney 1981)
and Hill et al (1982) observed the nymphs
clustering on plant leaves.
Riptortis linearis (F) 26 July 1995, J.
Santos leg, NEW RECORD, Coloane Is-
land. ex Phaseolus radiatus and P. vulgaris,
kidney bean. Wu (1935) reported it from
the Guangdong region of China as well as
India, Sri Lanka, Burma and Malaya.
Cydnidae
Adrisa magna (Uhler), black burrowing
bug. 12, 13 March; 3, 27 April 1994, ERE
leg, NEW RECORD, Friendship monu-
ment, Macau peninsula. It is also recorded
from Northeast India Burma and Hong
Kong.
Dinidoridae
Cyclopelta obscura (Lepeletier and Ser-
ville). 22 Sept. 1992 Ng Wai Man leg,
NEW RECORD, Coloane Island, ex Can-
avalia gladiata, sword bean and Cercis chi-
nensis, Chinese redbud. Distribution is In-
dia, Burma, Cambodia, Laos, Vietnam, In-
donesia, Malaysia, Philippines and China
(Durai 1987).
Megymenum inerme (Herrich-Schaeffer).
1 Aug. 1994, Ng Wai Man leg, Coloane
Island. Originally reported in Macau by
Kershaw (in Kirkaldy 1910). Also found in
India, Sri Lanka Vietnam, Thailand and
Guangzi, Guangdong, Hebei and Yunnan
provinces of China.
Gerridae
Aquarius paludum (EF). Summer 1993,
ERE and Leung-va leg, NEW RECORD,
Coloane Island in pond on reclaimed land
at Siac Pai Van; 21 Aug. 1996, Friendship
monument, Macau peninsula attracted to
lights. Chen and Andersen (1993) report it
from most provinces of China and it is be-
lieved to range across Eurasia from Britain
to Japan. These authors also reported an-
other species in Macau, namely Limnogon-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
us fossarum (FE) but we did not collect it in
this study.
Largidae
Physopelta gutta Burmeister. 28 Oct.
1992, Cheong Chi Kong leg, Coloane is-
land; 13 Feb.; 16, 27 March; 3, 6, 12 April
1994, ERE leg, Friendship Monument, Ma-
cau peninsula attracted to light. On the is-
land of Taipa and in neighboring Hong
Kong it has been noted (Easton 1992) feed-
ing on the seeds of the turn-in-the-wind
tree, Mallotus paniculatus, that mature dur-
ing the colder months of the year. Mating
was observed on the trunk of this tree and
all immature stages noted on the tree fo-
liage at this time. It is apparently wide-
spread in South East Asia found in Sri Lan-
ka, India, Burma, Vietnam, Bangladesh, In-
donesia, China (Guangdong, Hunan, Si-
chuan, Yunnan provinces), Tibet, Philippines,
Taiwan and east to Australia.
Lygaeidae
Horridipamera nietneri (Dohrn). 13 Aug.
1996, ERE leg, NEW RECORD, Friend-
ship Monument, Macau peninsula. In India,
Mukhopadhyay (1988) reported it from
grass and litter of Ficus species. It is also
found in the Guangdong Province of China
and reported as far southeast as Australia.
Metochus abbreviatus (Scott). 24 Aug.
1992, Cheong Pak Fai leg, Coloane Island;
10 Nov 1990, ERE leg, Taipa Island NEW
RECORD ex. Camellia oleifera, oil tea and
Vernicia montana, wood-oil tree. It is com-
monly attracted to lights. Distributed in In-
dia, China (Hunan, Guangxi, Jiangxi, Si-
chuan) Taiwan and Japan.
Metochus uniguttatus (Thunberg). 28
March 1996, ERE leg, Friendship monu-
ment, Macau peninsula. Easton (1992) re-
ported it from a building that had been il-
luminated at night on the island of Taipa.
There are also specimens collected from
Hong Kong at the Tai Lung farm experi-
mental station near Sheung Shui.
Paromius exiquus (Distant). 13 Aug.
1996, ERE leg, NEW RECORD, Friend-
VOLUME 99, NUMBER 3
ship monument, Macau peninsula attracted
to lights. Distribution includes India (Mu-
khopadhyay 1988).
Thunbergia sp. 27 July 1991, ERE leg,
Taipa island, attracted to lights on the uni-
versity campus.
Nabidae
Nabis stenoferus Hsiao. 15 Aug. 1992,
PWW leg, NEW RECORD Coloane island.
Also distributed in Hunan, Jilin, Shanxi,
Shandong and Yunnan provinces of China.
Notonectidae
Enithares biimpressa (Uhler). 6 June
1992, ERE leg, NEW RECORD, Coloane
island from small stream emerging from
mine shaft in ravine above Choec-van vil-
lage. It has been previously only reported
from Hong Kong (D. Polhemus, personal
communication).
Pentatomidae
Calliphara nobilus (L.), blue shield bug.
15 Aug. 1991, ERE leg, ex Ficus sp. near
entrance of mine shaft above Choec-van
village feeding on fruit. It was first reported
in Macau by Kirkaldy (1910). Distribution
includes Hong Kong, Borneo, Burma, Ma-
laysia, Java, Taiwan and the Philippines.
Cantao ocellatus (Thunberg). 5 Nov.
1994, ERE leg, Friendship monument, Ma-
cau peninsula. Easton (1991) reported nu-
merous individuals on the island of Taipa
resting and mating during winter months on
the fruit and among foliage of Mallotus pan-
iculatus (Euphorbiaceae). Eggs and nymphs
have not been observed in this study nor in
nearby Hong Kong where similar behaviour
has been reported but in India, parental care
has been noted (Ayyar 1920) and feeding
occurred on the tender leaves and fruit of
Trewia nudifolia (Euphorbiaceae), a plant
not found in this area. The insect further-
more in India was observed congregating on
the branches and so believed to be important
in the pollination of the moon tree, Macar-
anga roxburghii (Schuh and Slater 1995),
another plant species not believed to occur
Sy
locally or in Hong Kong. It is believed to be
distributed in Bhutan, Thailand, Burma,
Borneo, Java, Vietnam, Sumatra, Malaysia
and the Philippines as well as Taiwan and
southern Japan.
Chrysocoris stollii (Wolff). 7 July 1994.
Cheong Chi Kong leg, NEW RECORD,
Coloane Island, ex Glochidium and Schima
superba. Kershaw reported it first from Ma-
cau (in Kirkaldy 1910). It is also found in
Hong Kong, India, Java, Sumatra and the
Kwangsi province of China. Another spe-
cies Chrysocoris grandis (Thunberg), large
white shield bug was collected by J. C. Ker-
shaw 1910, but not found in the present
study.
Dalpada oculata (FE). 27 April 1993,
Cheong Pak Fai, leg, Coloane Island, ex
Cunninghamia lanceolata and Paulownia
fortunei. Kershaw reported it originally (in
Kirkaldy 1910) It is also found in the Hai-
nan province of China as well as in India,
Burma, Malaysia and Hong Kong.
Eocanthecona concinna (Walker). Hoff-
man (1935) listed it from both Hong Kong
and Macau but it was not observed in the
present study.
Eocanthecona furcellata (Wolff) grey
stink bug. 19 Aug. 1991, ERE leg, NEW
RECORD, Taipa island. A predaceous aso-
pine that feeds by inserting its stylets into
soft-bodied lepidopterous larvae. A mass
rearing technique has been developed to
control cutworms in Thailand (Napompeth
1992), and, in Hong Kong, it has been ob-
served feeding on the fall armyworm. Dis-
tribution includes India, Sri Lanka, Bang-
ladesh, Burma, Java, Taiwan, China to Ja-
pan (Thomas 1994).
Erthesina fullo (Thunberg), yellow spot-
ted stink bug. A common species locally
observed feeding on the trunks and stems
of Casuarina equisetifolia trees where mat-
ing and oviposition take place by cementing
two or more leaves together (Easton 1993).
First reported locally by Kershaw and Kir-
kaldy (1909c) who described the immature
stages and provided biological notes. In the
Guangzhou area of China, Hoffman (1930)
578
reported it feeding upon 30 species of trees
of which the tallow tree, Sapium sebiferum
was the first choice followed by the China-
berry, Melia azedarach and C. equisetifolia.
Its distribution includes India (Assam), Sri
Lanka, Burma (or Myanmar), Malaysia,
Java, Hong Kong, Taiwan and Japan.
Halyomorpha picus (F). 1 Aug. 1991,
ERE leg, Taipa Island. Hoffman (193 1a) re-
ported it as a pest of Vigna sesquipedalis
yard-long bean and Phaseolus lunatus, lima
beans near Guangzhou, China. We believe
Kershaw (in Kirkaldy 1910) was the first to
report it from Macau. It also occurs in Hong
Kong according to specimens at the Tai
Lung Experimental Station. Its distribution
includes India, Sri Lanka, Burma, Vietnam,
Malaysia and Taiwan as well as Guangdong
province of China.
Megarrhamphus hastatus (F). 10 May
1997, ERE leg, Taipa Island hiking trail, Ex
Miscanthus grass. Kirkaldy (1910) listed it
for southern China.
Melanophara dentata Haglund. Reported
in Macau by Kirkaldy (1910) but it has not
been collected in our study.
Nezara viridula (L.), the green vegetable
bug. One of the most common insects
found locally. Four colour varieties have
been observed in recent years. Of a total of
460 specimens collected from the lights of
the Friendship monument (Aug. 21—Sept. 8,
1996) an entirely green form comprized the
majority of specimens (85% of 460). A sec-
ond form characterized by a yellowish pro-
notum on a entirely green body accounted
for 13% while a 3rd form yellow on the
dorsal surface with green spots was the
least abundant (1.5%).
Only one individual has been observed
of the entirely yellow form which was char-
acterized by Chen (1980) and entered a
New Jersey Light trap on Taipa Island (1
Sept 1991). We believe Kirkaldy (1910)
was the first to report the apparently all
green form of this bug in Macau. Hosts of
the insect are numerous including Brassica
campestris, B. oberacea, B. pekinensis,
Oryza sativa and Azukia mungo.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Piezodorus hydneri (Gmelin), the soy-
bean stinkbug. 9 May 1996, ERE leg. Kir-
kaldy (1910) was first to record it locally.
In Hong Kong it is reported from Glycine
max or G. soja (Lee and Winney 1981).
Plautia fimbriata (EF), the string-bean
stinkbug. 8—10/tree 25 June 1996, ERE leg,
NEW RECORD, Taipa island, ex Clero-
dendrum fragrens, fragrant glorybower on
hiking trail feeding on flowers. In Hong
Kong it has been recorded feeding on Vigna
sesquipedalis, yard-long bean as well as in
the Guangdong region of China. It is also
found in India, Sri Lanka, Burma, Malaysia
and Japan.
Rhynchocoris humeralis (Thunberg), the
citrus shield bug. 19 Nov. 1994, ERE leg,
Friendship Monument, Macau peninsula
August 1996, Coloane island NEW REC-
ORD ex Citrus sinensis. Hoffman (1931b)
studied its life history in the Guangzhou
area of China and noted feeding on fruits
in all stages of development causing fruit
drop. It is found in India, Sri Lanka, Burma,
Thailand, Hong Kong and the Hainan,
Guangdong, Guangxi Fujian, Yunnan and
Sichuan Provinces of China as well as Tai-
wan and Laos.
Stollia guttigera (Thunberg). 14 Sept.
1993, PWW leg, Coloane Island. Kirkaldy
(1910) listed it earlier for Macau. Hosts in-
clude Amaranthus gracilus, Arachis hypo-
gaea, Avena fatua, wild oats, Celosia ar-
gentea, wild coxcomb, and Morus alba. It
is found in the Guangdong region of China.
Tetroda denticulifera Bergr, fork-headed
stinkbug. 14 June 1993, Taipa island; 13
Sept. 1991, ERE leg, Ka-Ho village, Co-
loane island, NEW RECORD. In Hong
Kong it has been recorded from Oryza sa-
tiva. Generic placement is considered to be
Tetrodias according to Ahmad and Kama-
luddin (1992).
Tolumnia latipes Walker. 27 April 1994,
Cheong Pak Fai leg, Coloane Island, ex Eu-
calyptus robusta, swamp mahogany. Ker-
shaw (in Kirkaldy 1910) may have recorded
it both in Hong Kong and in Macau as T.
latipes (Dallas). Distribution includes India,
VOLUME 99, NUMBER 3
Burma, Vietnam, Malaysia, Sumatra, Java,
Taiwan and the Hunan, Zhejiang, Jiangxi,
Hubei, Sichuan, Fujian, Guangxi, Yunnan
and Guangdong provinces of China.
Udonga spinidens Distant. 6 April 1995,
ERE leg, NEW RECORD, Friendship mon-
ument, Macau peninsula. It is widespread
in China found in the Fujian, Hubei, Hunan,
Guangxi, Guangdong, Shangxi and Yunnan
provinces as well as Vietnam and Japan.
Zicrona caerulea L. Kershaw and Kir-
kaldy (1909b) reported it in Macau feeding
on the chrysomelid beetle, Haltica coerulea
Olivier but it was not found in the present
study. In Hong Kong it has been recorded
from Oryza sativa (Lee and Winney 1981).
It is apparently widespread in China (Wu
1935) and found also in Burma (Myanmar),
Malaysia, India (Assam), Pakistan, Borneo,
Java, Sumatra, Vietnam and Taiwan.
Plataspidae
Brachyplatys subaeneus (Westwood). 1
June 1994, PWW leg, Coloane Island. Ker-
shaw (1910) reported it feeding on the Kud-
zu vine, Pueraria thunbergiana in Macau
and Hoffman (1931la) reported it on lima
beans in the Guangdong region of China. It
is also found in the Hainan province of Chi-
na as well as India, Thailand, Cambodia,
Malaysia, Burma, Borneo, Sumatra, Java,
the Philippines and Taiwan.
Coptosoma cribraria (F.) (= Megacopta
cribraria (F)). 8 Aug. 1994, Cheong Chi
Kong leg, Coloane Island. Kershaw (1910)
reported it in Macau. It is believed to feed
upon vetch, Phaseolus radiatus, and kidney
beans, P. vulgaris in the Guangdong region
of China. Also found in India, Sri Lanka,
Burma, Thailand, Java, Sumatra, Hong
Kong and Taiwan.
Coptosoma variegata Herrich-Schaeffer.
5 May 1994, Ip Tai leg, NEW RECORD,
Coloane Island. Also found in India, Sri
Lanka, Burma, Java, Borneo, Sumatra,
Thailand and the Philippines (Wu 1935).
Pyrrhocoridae
Dindymus rubiginosus sanguineus (F).
15 Nov., 9 Dec. 1990; 20 Jan. 1991 ERE
579
leg, Guia Hill on Macau peninsula. The first
records of this bug in Macau were by Ker-
shaw and Kirkaldy (1908a) who observed
nymphs and adults feeding on thin-shelled
gastropods, lepidopterous larvae, and he-
mipterans. Easton (1992) observed the bug
feeding on a worker ant near the base of
Pawlownia fortunei, Foxglove trees in a
forested area on the Guia hill. Feeding also
was noted on bugs of the same species at
this site that may have first been injured or
trampled by man. It is reported to occur in
India (Assam), Burma, Thailand as well as
Hong Kong (Wu, 1935).
Reduviidae
Ectomocoris apicimaculatus Distant. 7
Sept. 1995, ERE leg, NEW RECORD,
Friendship Monument, Macau peninsula. It
is also found in Hong Kong according to
records at Tai Lung Experimental Station,
Sheung Shui.
Ectomocoris atrox (Stal). 27 April 1993,
Ng Wai Man leg, Coloane Island; 14 Sept.
1995, ERE leg, Friendship Monument, Ma-
cau peninsula, NEW RECORD. Hua (1984)
recorded it from the Guangdong and Hai-
nan provinces of China and Maldonado Ca-
priles (1990) recorded it from Burma, Vi-
etnam, Malaysia, Indonesia, India as well
as China and the Philippines. This genus
(Schuh & Slater 1995) consists of species
that are ground-dwelling, and they are fast
moving bugs that can inflict a painful bite
to man.
Ectrychotes andreae (Thunberg). 2 April
1993, Tong Veng Va leg, NEW RECORD,
Coloane Island. The bugs in the subfamily
Ectrichodinae are believed to be obligate
predators of millipedes (Louis 1974) and
this species is found in the Guangdong and
Hainan provinces of China (Hua 1984) as
well as Taiwan, Korea, Japan and the In-
dochinese area (Maldonado Capriles 1990).
Oncocephalus impudicus Reuter. 14
Sept. 1995, 15 Feb. 1996, Friendship Mon-
ument, Macau peninsula; 30 Sept 1990,
Univ. E. Asia, Taipa Island, ERE leg, NEW
RECORD. Its distribution includes Borneo,
580
Java and Sumatra in Indonesia, Sri Lanka
and the Guangzhou area (Guangdong) and
Hainan provinces of China.
Polididus armatissimus Stal. 9, 14 Sept,
1995, ERE leg, NEW RECORD, Friend-
ship Monument, Macau peninsula. Its dis-
tribution includes India, Sri Lanka, Burma
(Myanmar) Vietnam, Malaysia and the Fu-
jian, Jiangxi, Hubei, Zhejiang, Guangxi,
Guangdong and Hainan provinces of China.
Scadra costalis (Lethierry). 4 June 1996,
ERE leg, NEW RECORD, Coloane island
crossing road on: premises of the Macau
Golf and Country Club near Hac-sa village.
This genus is also believed to feed upon
millipedes which are quite common locally.
Sycanus crocevittatus Dohrn. 12 Aug.
1996, ERE leg, on causeway between Taipa
and Coloane islands. Kershaw (1909) dis-
cussed oviposition and development of this
bug in South China (probably included
Hong Kong and Macau) and Hill et al.
(1982) in Hong Kong discussed it feeding
when a nymph on snails. It is found in India
and Burma as well as islands off the coast
of Guangdong province and on Hainan is-
land (Hua 1984).
Triatoma rubrofasciata (DeGeer). 23
Aug. 1995, ERE leg, Taipa island attracted
to lights on the University campus. Easton
(1992) noted several individuals under
rocks on the ground surface near a rodent
harborage. Usinger (1944) pointed out that
even though its distribution is currently tro-
picopolitan it is considered an old world
species and probably Indian in origin. It oc-
curs in Hong Kong as well as Hainan is-
land.
Tribelocephala walkeri China. 15 July
1972. Chan Chi Man leg, NEW RECORD,
Coloane Island. It occurs in Hong Kong
(Maldonado Capriles 1990) and Hua (1984)
reported it from islands off the coast of the
Guangdong province.
Tessaratomidae
Tessaratoma papillosa (Drury), litchi
stinkbug. 11 Nov. 1992, Ng Wai Man leg,
Coloane Island; 5 Sept. 1994, ERE leg, Tai-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
pa island. Kershaw and Muir (1909) were
probably the first to record it from Macau.
It is common locally and in Hong Kong on
both longan (Euphoria longan) and lychee,
Litchi chinensis fruit trees where ever they
occur. Considered a pest of fruit trees, bi-
ological control has been achieved by the
mass rearing and release of an egg parasite,
Anastatus japonicus Ashmead (Eupelmi-
dae: Hymenoptera) in Hong Kong and in
the Guangdong province of China as well
as in Thailand (Napompeth 1992). It is also
found in India, Sri Lanka, Burma, Thailand,
Java, Sumatra and the Philippines.
Tingidae
Stephanitis pyrioides (Scott). azalea lace
bug. 25 Mar. 1994, PWW leg, NEW REC-
ORD, Coloane Island. In Hong Kong this
species has been collected from Rhododen-
dron indicum (Lee & Winney 1981), and
there are records from Purple azaleae, R.
pulchrum, in the Tai Lung farm entomology
collection.
ACKNOWLEDGMENTS
The authors are much appreciative to Mr.
Clive S. K. Lau, Plant Protection officer,
Agriculture and Fisheries, Hong Kong Gov-
ernment, for allowing one of us (ERE) ac-
cess to their insect collection on the Tai
Lung Experimental Station near Sheung
Shui, New Territories, Mr. Lau also ar-
ranged with The Natural History Museum,
London to have identifications carried out,
such as Thunbergia sp. of Lygaeidae that
was determined by G. M. Stonedahl. We are
indebted to Mr. Anténio Julio Emerenciano
Estacio, Head of Agrarian services, Island
government, for permission to use their fa-
cilities at Siac Pai Van. Species determina-
tions of the gerrid and notonectid water
bugs were made by Dr. Dan Polhemus,
Smithsonian Institution, Washington, D.C.;
identifications of pentatomids, in part, by
Prof. Ghen Zhenyao, Research Institute of
Entomology, Zhongshan University,
Guangzhou, China, and identification of the
Lygaeidae were made, in part, by Dr. James
VOLUME 99, NUMBER 3
A. Slater of the University of Connecticut,
Storrs.
LITERATURE CITED
Ahmad I. and S. Kamaluddin. 1992. New generic sta-
tus of a rice-feeding Tetrodine subgenus, Tetro-
dias Kirkaldy and redescription of Tetroda Amyot
and Serville and their cladistic analysis. Pakistan
Journal of Zoology 24(2) 123-27.
Ayyar, T. V.R. 1920. Notes on the life-history of Can-
tao ocellatus, Th. Report of the Proceedings of the
Third Entomologial Meeting, Pusa, III: 910-914.
Calcutta Superintendent Government Printing, In-
dia.
Chen, EF 1980. A new form of Nezara viridula (Lin-
naeus). Entomotaxonomia 2(1) 31-32. (In Chi-
nese.)
Chen, P-P. and N. M. Andersen. 1993. A Checklist of
Gerromorpha from China (Hemiptera). Chinese
Journal of Entomology 13: 69-75.
Dudgeon, D. and R. Corlett. 1994. Hills and Streams,
an ecology of Hong Kong. Hong Kong University
Press. 234 pp.
Durai, P. S. S. 1987. A revision of the Dinidoridae of
the World (Heteroptera:Pentatomoidea). Oriental
Insects 21: 163-360.
Dolling, W. R. 1978. A revision of the Oriental pod
bugs of the tribe Clavigrallini (Hemiptera:Corei-
dae). Bulletin of the British Museum (Natural His-
tory), Entomology. 36(6) 281-321.
Easton, E. R. 1991. Annotated list of the insects of
Macau observed during 1989. Entomological
News 102(2): 105-11.
Easton, E. R. 1992. 1990 additions to the annotated
list of the insects of Macau. Entomological News
103(1): 30-36.
Easton, E. R. 1993. The Insects of Macau. University
of Macau publication Centre. 58 pp.
Hill, D. S., P. Hore, and I. W. B. Thornton. 1982. In-
sects of Hong Kong. Hong Kong University Press.
503 pp.
Hoffman, W. E. 1930. The food habits of Erthesina
fullo (Thunb.) Lingnan Science Journal 9: 139—
142.
Hoffman, W. E. 193la. Notes on Hemiptera and Ho-
moptera at Canton Kwangtung province, Southern
China, 1924-1929. Bureau of Entomology. Unit-
ed States Department of Agriculture Insect Pest
Survey Bulletin 11: 138-151.
Hoffman, W. E. 1931b. The life history of Rhyncho-
coris humeralis Thunb. (Hemiptera:Pentatomi-
dae). Lingnan Science Journal 7: 817-820.
Hoffman, W. E. 1933. Additional data on the life his-
tory of Lethocerus indicus (Hemiptera:Belosto-
matidae). Lingnan Science Journal 12: 595-601.
Hoffman, W. E. 1935. An abridged Catalogue of cer-
tain Scutelleroidea (Plataspidae, Scutelleridae, and
Pentatomidae) of China, Chosen, Indo-China and
581
Taiwan. Lingnan University Science Bulletin
Number 7, 294 pp.
Hua Lizhong. 1984. A Check list of reduviid bugs of
Guangdong Province (Hemiptera:Reduviidae). In-
stitute of Entomology, Zhongshan University,
Guangzhou, China. 18 pp. (In Chinese.)
Kershaw, J. C. W. 1909. On the metamorphoses and
anatomy of the reduviid bug, Sycanus croceovit-
tatus Dohrn. Annales de la Societe Entomologique
de Belgique 53: 241—249.
Kershaw, J. C. W. 1910. On the metamorphoses of
two coptosomine hemiptera from Macao. Annales
de la Société Entomologique de Belgique 54: 69—
73:
Kershaw, J. C. W. and G. W. Kirkaldy. 1908a. Bio-
logical notes on Oriental Hemiptera, No. 1. Jour-
nal of the Bombay Natural History Society 18:
596-598.
Kershaw, J. C. W. and G. W. Kirkaldy. 1908b. On the
metamorphoses of two hemiptera-heteroptera
from southern China. Transactions of the Ento-
mological Society of London, pp 59-62.
Kershaw, J. C. W. and G. W. Kirkaldy. 1909a. Bio-
logical notes on Oriental hemiptera II. Journal of
the Bombay Natural History Society 19: 177-178.
Kershaw, J. C. W. and G. W. Kirkaldy. 1909b. Bio-
logical notes on oriental Hemiptera III. Journal of
the Bombay Natural History Society 19: 333-336.
Kershaw, J. C. W. and G. W. Kirkaldy. 1909c. Bio-
logical notes on oriental hemiptera. IV. Journal of
the Bombay Natural History Society 19: 571-573.
Kershaw, J. C. W. and E Muir. 1909. Life history of
Tessaratoma papillosa Thunberg with notes on
the stridulating organ and stink glands by Fred-
erick Muir. Transactions of the Entomological So-
ciety of London, pp 253-258.
Kirkaldy, G. W. 1910. A list of the Hemiptera of Ori-
ental China, 2. Annales de la Société Entomolo-
gique de Belgique 54: 103-112.
Lee, L. H. Y. and R. Winney. 1981. Checklist of Ag-
ricultural insects of Hong Kong 1981. Agriculture
and Fisheries Department Bulletin No. 2, 164 pp.
Louis, D. 1974. Biology of Reduviidae of cocoa farms
in Ghana. American Midland Naturalist 91: 68—
89.
Maldonado Capriles, J. 1990. Systematic catalogue of
the Reduviidae of the world (Insecta:Heteroptera).
Caribbean Journal of Science (special edition),
694 pp.
Mukhopadhyay, A. 1988. Taxonomic study of Ly-
gaeidae (Heteroptera:Insecta) from West Bengal
(India). Records of the Zoological survey of India,
Miscellaneous Publication Occasional paper No.
107, 72 pp., 12 plates.
O’Shea, R. and C. W. Schaefer. 1980. A generic re-
vision of the Asian and Australian Mictini (Het-
eroptera:Coreidae). Oriental Insects 14: 221—251.
Napompeth, B. 1992. Brief survey of Biological Con-
582 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
trol activities in Thailand, pp. 51-68. In Hirose, Thomas, D. B. 1994. Taxonomic synopsis of the Old
Y. ed., Biological control in South and East Asia. World asopine genera (Heteroptera:Pentatomidae).
Kyushu University Press, Fukuoka, Japan. Insecta Mundi 8 (3-4): 145-212.
Sands, D. PR A. 1977. The Biology and ecology of | Usinger, R. L. 1944. The Triatominae of North and
Leptocorisa in Papua New Guinea. Research Bul- Central America and the West Indies and their
letin No. 18, Department of Primary Industry, Port public health significance. United States Public
Moresby. 104 pp. Health Service Bulletin 288, 83 pp. 11 plates.
Schuh, R. T. and J. A. Slater. 1995. True bugs of the Wu, C. E 1935. Catalogus Insectorum Sinensium
world (Hemiptera:Heteroptera) classification and (Catalogue of Chinese Insects). Volume II. Fan
natural history. Comstock Publishing, Cornell Memorial Institute of Biology, Peiping, China.
University Press, Ithaca. 336 pp. 634 pp.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 583-584
NOTE
Acoptus suturalis LeConte (Coleoptera: Curculionidae: Zygopinae),
a Potential Vector of the Chestnut Blight Fungus,
Cryphonectria parasitica (Murrill) Barr, in the Eastern United States
American chestnuts trees, Castanea den-
tata (Marshall) Borkhausen, were once a
widespread and important component of
forests throughout the eastern United
States. Following the introduction of the
chestnut blight fungus, Cryphonectria par-
asitica (Murrill) Barr, into this country ear-
ly in the century there was a rapid and dra-
matic decline in the abundance of these
trees (Anagnostakis 1994. Advances in Bo-
tanical Research 21: 125-145). In an at-
tempt to reestablish this species in forests
and orchards there has been an exhaustive
search for blight-resistant tree strains by
cross-breeding with other species of Cas-
tanea (Anagnostakis 1994). More recent
studies suggest that infected chestnuts in-
oculated with certain hypovirulent strains of
the fungus are less susceptible to severe
canker-induced damage (Anagnostakis
1994). Hypovirulence, caused by infection
of the fungus with a double-stranded RNA
virus or hypovirus, reduces the fungus dam-
age to trees. Transgenic fungal strains have
been produced that have a cDNA copy of
the Hypovirus genome integrated into the
genome of the blight fungus (Choi and
Nuss 1992. Science 257: 800-803). These
hypovirulent strains have the potential to
effect a stable biological control of chestnut
blight.
Dissemination of these hypovirulent
strains outside the area of test plots will de-
pend upon naturally occurring dispersal
agents. Since insects, and particularly bee-
tles, are known to harbor hypovirulent
strains (Russin et al. 1984. Journal of Eco-
nomic Entomology 77: 838-846) we were
interested to find large numbers of the wee-
vil Acoptus suturalis LeConte (Coleoptera:
Curculionidae: Zygopinae) feeding upon
chestnut cankers in an experimental plot
(150 m X 80 m) in the Housatonic State
Forest (Sharon township, Litchfield county,
73 22 30W X 41 52 40N, elevation 395 m)
in northwestern Connecticut. In this forest
test plot, pairs of American chestnut trees,
AB to WX, were used in a study of the
effect of virulent strains of the blight fun-
gus. The first tree of each pair was treated
with a transgenic hypovirulent strain, with
resistance to hygromycin as an additional
marker.
The vast majority of curculionids feed
upon the living tissue of diverse structures
of plants, although a considerable number
are associated with decaying wood or fungi.
Little is known about the feeding associa-
tions for zygopine weevils, although adults
are often found in the vicinity of rotting
wood, so the presumption is that the larvae
develop in this habitat (Hespenheide 1995.
Memoirs of the Entomological Society of
Washington 14: 145-154). Lyal (1986.
Journal of Natural History 20: 789-798)
observed species of mecopine and metial-
mine Zygopinae ovipositing in the bark of
fallen trees in southeastern Asia. The zyg-
opine weevil Acoptus LeConte, a monotyp-
ic genus, is represented by A. suturalis and
is widely distributed in eastern North
America. Adults of this species have been
taken from Quercus sp., Cercis canadensis
L., Carya sp., and Platanus occidentalis L.
(Sleeper 1963. Bulletin of the Southern
California Academy of Sciences 62(4):
209-220), and from chestnut (Russin et al.
1984). Additionally, Chittenden (1890. En-
tomologica Americana 6: 167—172) report-
ed Acoptus suturalis adults and supposedly
conspecific larvae living in the dead wood
of beech trees (Fagus sp.).
584
lait, It
blight cankers infected with a hypovirulent transgenic
strain of Cryphonectria parasitica.
Acoptus suturalis feeding on chestnut
Acoptus suturalis weevils were observed
in Connecticut grazing on the fungal stro-
mata in the Cryphonectria parasitica can-
kers on American chestnut trees (Fig. 1) in
the test plot which is surrounded by oak-
chestnut forest. These weevels were not ob-
served on other woody plants in the plot.
Trees in the overstory are Quercus coccinea
Muench., Acer rubrum L. and Populus sp.,
while woody plants in the understory in-
clude Carya glabra (Miller) Sweet, Quer-
cus prinus L., Acer pennsylvanicum L.,
Hamamelis virginiana L. and an Ulmus sp.
One of us (SLA) collected 36 weevils (al-
though many others were observed in the
test plot), surface sterilized them with a
10% bleach solution, and squashed and
spread them on water agar. Of 280 fungal
isolates, 13 were hygromycin resistant, in-
dicating that they contain transgenic DNA.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
These observations suggest that Acoptus
weevils may play a role in controlling
chestnut blight in eastern North America
since the abundance of these weevils in test
plots suggests their potential for spreading
a transgenic hypovirulent strain which may
eventually help to control chestnut blight.
Acknowledgments.—We are indebted to
Henry A. Hespenheide (University of Cal-
ifornia, Los Angeles) for information about
zygopine weevil biology and some relevant
literature. We thank Horace R. Burke (Tex-
as A & M University), Joseph V. McHugh
(University of Georgia), P. Sletten and J.
Shepard (The Connecticut Agricultural Ex-
perimental Station, New Haven), and Allen
L. Norrbom (Systematic Entomology Lab-
oratory) for reviewing this manuscript. This
research was supported, in part, with funds
from the Cooperative State Research, Edu-
cation, and Extension Service, U. S. De-
partment of Agriculture, under agreement
No. 95-37312-1638.
James Pakaluk, Systematic Entomology
Laboratory, PSI, Agricultural Research
Service, U.S. Department of Agriculture,
c/o National Museum of Natural History,
MRC-168, Washington, DC 20560, U.S.A.;
Sandra L. Anagnostakis, The Connecticut
Agricultural Experiment Station, Box 1106,
New Haven, CT 06504, U.S.A.
VOLUME 99, NUMBER 3
585
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 585-586
NOTE
Parasa indetermina (Boisduval) (Lepidoptera: Limacodidae), a new host for Systropus
macer Loew (Diptera: Bombyliidae)
This note adds a new species to the
published limacodid hosts of Systropus
macer Loew (Diptera: Bombyliidae). Host
literature for worldwide Systropus was re-
viewed by Adams and Yanega (1991,
Journal of the Kansas Entomological So-
ciety 64:443—444). All North American
records have been restricted to the eastern
U.S. and are reported only for S. macer.
Forty-nine species of Limacodidae are
known from North America north of Mex-
ico, accounting for synonymies from Ep-
stein and Becker (1993, Revista brasileira
de Zoologia 10:289—319) and a newly es-
tablished species in southern Texas (Fer-
guson and Knudson 1987, Journal of the
Lepidopterists’ Society 40:353-—355). Of
these species, only four have been report-
ed as hosts for Systropus: Euclea delphi-
nii (Boisduval), Adoneta spinuloides
(Herrich-Schaffer), Prolimacodes badia
(Hiibner), Lithacodes fasciola (Herrich-
Schiffer), and an unidentified larva of
Apoda Haworth (Adams and Yanega
1991). There are no additional unpub-
lished host records for Systropus in the
collection of the National Museum of Nat-
ural History, Smithsonian Institution
(USNM), presently on loan to the Bishop
Museum (Neal Evenhuis personal com-
munication).
Larvae of Parasa indetermina (Boisdu-
val) and one larva of Euclea delphinii (Bois-
duval) were found feeding on bayberry near
Higbee Beach, Cape May, New Jersey on
Sept. 23—24, 1995. All spun cocoons over
the following several weeks. Adults of the
non-parasitized cocoons all emerged in June
1996. The larva of S. macer inside the co-
coon of E. delphinii was discovered in early
June 1996 and was kept in a 2 dram vial. It
did not pupate until 10 July 1996. It emerged
along with the other individuals that parasit-
ized P. indetermina between 10—15 August
1996. One individual per cocoon emerged in
characteristic fashion, by pushing open the
lid of the cocoon, as occurs for the limacod-
ids. Five out of 17 cocoons of P. indeter-
mina produced Systropus adults.
Additional New World species of Par-
asa that are hosts of Systropus sp. are P.
wellesca Dyar and P. cuernavaca Dyar
from Guanacaste, Costa Rica (D.H. Jan-
zen and W. Hallwachs Lepidoptera rearing
record database). There are no records of
Parasa parasitized by Systropus in a re-
view of bombyliid parasitoids of South-
east Asian limacodids by Greathead
(1987, pp. 195-196. In Cock, Godfray
and Holloway, eds., Slug and Nettle Cat-
erpillars. CAB International). However,
several species in the African limacodid
genus Latoia (as Parasa) have been re-
ported as hosts (Bowden 1967, Journal of
the Entomological Society of Southern
Africa 30:126—173).
Published limacodid hosts of North
American and Asian Systropus consist of
both smooth (= gelatine) and spiny type
larvae; examples of smooth larval hosts in-
clude Prolimacodes Schaus, Lithacodes
Packard and Chalcocelis Hampson (Adams
and Yanega 1991; Greathead 1987). Re-
cords from Mexico south to Argentina are
exclusively from the spiny genera Acharia
(=Sibine) Hiibner and Miresa Walker (lit-
erature reviewed by Adams and Yanega
1991) and the two Parasa species above.
This probably reflects a collecting bias for
brightly colored spiny larvae, which tend to
586
be agricultural pests, compared to the cryp-
tic smooth larvae.
I thank Jane Ruffin (Rosemont, PA) for
discovering and showing me the population
of Parasa indetermina and Neal Evenhuis
(Bishop Museum) for determining the spec-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
imens of S. macer, providing helpful bib-
liographic information and searching the
USNM collection for host information.
Marc E. Epstein, Department of Ento-
mology, MRC 105, Smithsonian Institu-
tion, Washington, D.C. 20560, U.S.A.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 586
NOTE
Replacement Names for Western Hemisphere Genera of Doryctinae
(Hymenoptera: Braconidae)
Recently I published descriptions of sev-
eral new genera of Doryctinae from the West-
erm Hemisphere (Marsh 1993. Contributions
of the American Entomological Institute
28(1): 1-58). Unfortunately, three of these
generic names have been previously used and
must be replaced with new names. Therefore,
I propose the following new generic names.
Cyphodoryctes Marsh, new name. Re-
placement name for Cyrtonion Marsh 1993,
not Cyrtonion Hansen 1989 (see Hansen
1991. Kongelige Danske Videnskabernes
Selskab Biologiske Skrifter 40: 1-367). The
original name Cyrtonion was derived from
the Greek kyrton meaning humpbacked in
reference to the mesonotum which anteri-
orly is swollen and strongly declivous. The
replacement ‘name is derived from the
Greek kyphos also meaning humpbacked.
Donquickeia Marsh, new name. Re-
placement name for Quickia Marsh 1993,
not Quickia Odhner 1950 (see Vaught 1989.
A classification of the living Mollusca.
American Malacologists, Inc.).
Whitfieldiellus Marsh, new name. Re-
placement name for Whitfieldia Marsh
1993, not Whitfieldia Davidson 1882 (see
Moore, ed., 1965. Treatise on invertebrate
paleontology. Part H, Brachiopoda. Vols. 1
and 2. The Geological Society of America
and the University of Kansas).
Paul M. Marsh, P.O. Box 384, North
Newton, KS 67117, U.S.A.
VOLUME 99, NUMBER 3
587
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 587
NOTE
Gender Correction for a Louse (Phthiraptera: Menoponidae) from Cormorants
Eichler (1950, Rivista di Parassitologia
11: 106) described Piagetiella caputincisa
(Phthiraptera: Menoponidae) as a new spe-
cies, with the type host being the blue-eyed
cormorant, Phalacrocorax atriceps King.
The agreement of the specific with the ge-
neric name is obvious, as both are feminine.
Subsequent workers have been consistent in
perpetuating the spelling given by Eichler.
These include Hopkins and Clay (1952,
British Museum (Natural History): 292) in
their check list of the genera and species of
chewing lice, Price (1970, Canadian Ento-
mologist 102: 401) in his revision of the
genus Piagetiella, Clay and Moreby (1970,
Pacific Insects Monograph 23: 219) in a
listing of lice of subantarctic islands, and
Robinson (1984, British Museum (Natural
History) Publication Number B2 0955: 31)
in a checklist of insects of the Falkland Is-
lands. To the best of our knowledge, all
published references to this species have
spelled it with the species in the feminine
gender.
The late George C. Steyskal, in his ef-
forts to put Latin ignorami on the right path,
wrote to explain to us the error of our ways
in considering “‘caputincisa”’ as the correct
form. According to him, “‘caputincisa’”’ rep-
resents a noun-phrase in apposition and the
adjective “‘incisa”’ refers to “‘caput’’, which
is a neuter noun. Therefore, the correct
spelling for this scientific name must be Pi-
agetiella caputincisum. Rather than simply
change this spelling in our future references
to this species, we thought it prudent to call
attention as to why we are taking this action
and remove any doubt that we may have
committed a typographical error in so do-
ing.
Roger D. Price, 4622 Kinkead Ave., Fort
Smith, AR, U.S.A. 72903; Ricardo L. Palma,
Museum of New Zealand, P.O. Box 467,
Wellington, New Zealand.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 588-590
ROBERT TRAUB (1916—1996): AN APPRECIATION
Historians are fond of reminding us that
the nineteenth century effectively began
with Wellington’s victory at Waterloo in
1815 and ended with the guns of August
1914. But do they know that the twentieth
century also has ended off schedule? At
least for medical entomology, the curtain
fell on 21 December of last year, when we
lost Dr. Robert Traub, Colonel, U.S. Army
(Retired), the greatest authority on fleas and
flea-borne diseases who ever lived and one
of the most influential medical entomolo-
gists of this—or any—century. In months
ahead, Dr. Traub’s panegyrists will be le-
gion, and formal obituaries may be expect-
ed in scores of scientific journals and so-
cietal newsletters worldwide. This is not
one of them. It is, rather, a reminiscence, an
anecdotal adieu to both a preeminent intel-
lect and a charming personality.
It has been said that success in life is
largely a matter of luck, in which case Rob-
ert Traub was lucky from the cradle, for he
was born on 26 October 1916 in the Bor-
ough of New York (Manhattan), the cultural
center of a city universally renowned for its
tolerance and generosity. Unfettered by pro-
vincialism and imbued from childhood with
a passion for natural history, Traub soon se-
cured what was, for his generation, the
Holy Grail of academe: admission to the
City College of New York, which, by virtue
of being tuition-free, attracted the cream of
America’s youth throughout the Great De-
pression. Success at City College in 1938
(B.S. in biology, cum laude) was Traub’s
entrée to the Ivy League; the following
year, Cornell University awarded him an
M.S. in his chosen métier of medical en-
tomology. But just three months after the
commencement ceremonies, Hitler un-
leashed 52 divisions against Poland, and en-
tomologists everywhere felt the blow.
Paradoxically, the massive deployment
of Allied forces during World War II her-
alded medical entomology’s golden age,
particularly in tropical Asia. Dispatched as
a young Army officer to the China-Burma-
India Theater, Traub’s attention soon fo-
cused on a mysterious illness initially
dubbed ‘“‘CBI fever’ that turned out to be
scrub typhus (chigger-borne rickettsiosis,
tsutsugamushi disease), an often fatal infec-
tion that was feared more than malaria be-
cause its etiology was unknown. Working
at the United States of America Typhus
Commission laboratory in Myitkyina,
North Burma, and, after the war, at the U.S.
Army Medical Research Unit, on the
grounds of the Institute for Medical Re-
search, Kuala Lumpur, Malaya, Robert
Traub and his American and British collab-
orators clarified the intricate ecology of this
disease and ultimately defeated it with the
potent new antibiotic chloromycetin (chlor-
amphenicol). These epochal advances in ty-
phus research earned Traub’s team a nom-
ination for the Nobel Prize in 1948.
VOLUME 99, NUMBER 3
Acarologists will always claim Robert
Traub as one of their own. Yet, even while
he immersed himself in mite studies (pub-
lishing over his lifetime 66 papers on chig-
gers and chigger-borne rickettsiosis, includ-
ing descriptions of 124 new chigger spe-
cies), Traub was cultivating his lifelong en-
thusiasm for what his friend and mentor
Karl Jordan (1861—1959) called ‘‘those jol-
ly insects’’: fleas. Indeed, four new species
of North American fleas were the subject of
Traub’s first scientific paper, published by
the Field Museum in 1944, while its author
was away pursuing acarines in the Burmese
jungles (fleas will also be Traub’s swan
song, in the form of a co-authored chapter
in a new textbook of medical and veterinary
entomology slated to appear next year).
Tellingly, 31 of Traub’s 91 papers on fleas
and flea-borne diseases were written in his
spare time during a military career (1942—
1962) that kept him otherwise occupied.
And what occupations! While with the
Army, Traub completed his Ph.D. at the
University of Illinois (1947), in the com-
pany of his confrere Harry Hoogstraal
(1917-1986), whose name will forever be
a mantra in medical acarology. After the
Army, Traub accepted a professorship in
the Department of Microbiology, Universi-
ty of Maryland School of Medicine, Balti-
more, subsequently leading or participating
in 14 research expeditions on four conti-
nents. During both careers, his published
contributions, many of book length, on
fleas, mites, mosquitoes, leeches, even am-
ebiasis, as well as on a vast range of host
and reservoir vertebrates, defined the state
of the art in parasitology. And all the while,
he was the exemplar of domesticity—proof
that professional achievement need not
(must not!) eclipse familial devotion.
Science has seldom had a happier expo-
nent than Robert Traub, whose charm was
disarming. To this day, throughout the
Third World, even the least of his former
technicians remember him with unbridled,
often tearful affection. Though a military
man, tough ‘‘three-gun Traub” never
589
stooped to the language of the gutter;
though honored the world over, he never
lost the common touch. His humor could
leave an audience in stitches, as when he
would explain the derivation of the specific
epithet fujigmo (no, it’s not a Japanese sur-
name!), applied to a new chigger by his sar-
donic associates Cornelius B. Philip and
Henry Shepard Fuller (1950, Journal of
Parasitology 40: 50-57). One of his favor-
ite recollections was of taking a company
of soldiers on an extended field exercise
through waist-deep swamps in Southeast
Asia. After several days, Traub noticed that
the company “‘ration”’ of condoms was run-
ning low, but his men at least seemed to
have overcome their fear of the swamps’
purportedly urethraphilous leeches.
Traub often applied the evocative de-
scriptor “‘Renaissance man” to the lumi-
naries in his life, among them the peerless
medical ecologist J. Ralph Audy (1914-
1974) of “Imphal Circus” days (Audy’s
Red Mites and Typhus, published in 1968
by the Athlone Press, University of Lon-
don, may well be the most engaging acar-
ological essay ever written) and, of course,
Harry Hoogstraal. But it took one to know
one. Traub’s interests, like his profession,
embraced the world. His erudition was im-
mediately apparent in such spheres as his-
tory and classical music, but he was also a
connoisseur of fine wines, exotic foods, and
East Asian objets trouvés, especially those
crafted of jade, ivory, or rare woods; of
these latter, he amassed invaluable personal
collections. Touchingly, he retained a child-
like fascination with the films of “Stan”
Laurel and Oliver Hardy, the first great
comedy team of the “‘talkies’’; a poster of
the perennially befuddled “‘boys”’ was a fix-
ture in his basement laboratory.
The last decades of Traub’s life were vir-
tual excerpts from the Book of Job. Among
his afflictions: pernicious diabetes, recur-
ring cancers and, most painful to recall, the
loss in 1989 of his gifted daughter Jean-
nette—all against the backdrop of the oft-
untimely passing of his former comrades-
590
in-arms. Yet, in the 20 years we knew him,
not once did we hear him complain. Sus-
tained by Renée, his wife and foil of 57
years, and their accomplished son Roger,
Traub’s tireless wit prevailed at death’s very
door, as in an exchange overheard at the
National Naval Medical Center, in Bethes-
da, Maryland. Asked by a Dr. Watson to
describe his latest problem, Traub replied:
‘‘Alimentary, my dear Watson.”
A week before Traub’s death, we made a
final pilgrimage to his bedside. We found
him incoherent, already beyond reach of
our farewells. Had he been able to under-
stand us, what could we have said to this
man, who all his adult life was contemp-
tuous of his own feelings but ever empa-
thetic with others? Since adolescence, one
of us (RGR) has kept on his night table the
thoughts of the Roman Emperor and Stoic
philosopher Marcus Aurelius Antoninus
(A.D. 121-180), as translated by the great
John Jackson for Oxford University Press
in 1906 and reprinted down to this day. An-
ticipating by more than a millennium the
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Florentine triumph that gave us Robert
Traub, Antoninus offers these words of
parting (XII: 36): “‘Friend, thou hast been
a citizen in this great city, and what matters
it whether for five years or three? The law
is the same for us all. Where is the hard-
ship, then, if it be no tyrant’s stroke, no un-
just judge, that sends thee into exile, but the
same Nature that brought thee hither, even
as the master of the show dismisses the
mummer that he put on the stage?... For
He decrees it shall end, who was once the
author of thy existence, and now of thy dis-
solution .... Then depart in peace with all
men, for He who bids thee go is at peace
with thee.”
Richard G. Robbins, Armed Forces Pest
Management Board, Walter Reed Army
Medical Center, Forest Glen Section,
Washington, DC 20307-5001, U.S.A.;
Ralph P. Eckerlin, Division of Natural Sci-
ences and Mathematics, Northern Virginia
Community College, 8333 Little River
Turnpike, Annandale, VA 22003-3796,
U.S.A.
PROC. ENTOMOL. SOC. WASH.
99(3), 1997, pp. 591-592
INSTRUCTION FOR AUTHORS
General Policy.—Articles for publica-
tion are ordinarily accepted only from
members of the Society and must be in En-
glish. A summary in French, German,
Spanish, or Russian is acceptable. Such a
summary should be placed immediately af-
ter the English abstract and may be a trans-
lation of that abstract. Manuscripts should
not exceed 15 printed pages (about two and
one-half double-spaced typewritten pages
equal one page). Manuscripts are peer-re-
viewed before acceptance. Final acceptance
is the responsibility of the Editor. Articles
are published in the order received, not the
date of acceptance. Immediate publication
is available with payment of full page
charges. Notes, book reviews, and obituar-
ies are published as space is available.
Manuscript preparation.—Submit three
printed copies, including copies of illustra-
tions, to the Editor. It is not necessary to
send original illustrations until requested af-
ter the manuscript has been accepted. After
acceptance, the final version may be sub-
mitted on diskette (DOS or Macintosh,
preferably WordPerfect or Microsoft Word;
no ASCII files), but a hard copy is also re-
quired. Use letter-sized paper. The entire
manuscript should be double spaced. Allow
adequate (1—-1%”) margins on all sides.
Words to be italicized may be indicated in
italic form or underlined. Manuscript con-
tents should be in the following sequence
and items 1—6 should begin a new page:
1. Title page with title, author(s), and au-
thor(s) byline. In upper left, give
name, address, phone, fax, and e-mail
of the author to whom all correspon-
dence and proofs are to be sent. Titles
must include order and family place-
IMEM EO ay Bice 5s os (Hymenoptera:
Xyelidae). Taxa of the genus and spe-
cies groups levels must have the au-
thority name.
2. Abstract; key words
3. Text of paper; acknowledgments at
end
. Literature Cited (see below)
. Figure legends
. Appendix; footnotes (if any)
. Copies of tables; legends for tables
are placed at the top of each table.
8. Copies of figures.
NANA
Illustrations.—There is no extra charge
for line drawings or half tones. Authors
should plan illustrations for reduction to the
dimensions of the printed page and allow
room for legends at the bottom. Do not
make plates larger than 14 X 18”. Individ-
ual figures should be mounted on suitable
board, and photographs (glossy only)
should be trimmed, grouped together, and
abutted when mounted; the printer will in-
sert hairlines. Figure numbers are prefera-
bly placed at the lower right of each figure.
Arrange plates so there is as little wasted
space as possible. Include title of paper, au-
thor(s), name and address, and illustration
numbers on the back of each plate. Figures
should be numbered consecutively and usu-
ally in the sequence in which they are re-
ferred to im the text. Reference mi text
should be as “Fig. 15> “Figs: 2; 3; la-
ble 1.’’ See recent Proceedings for style of
figure legends.
Literature Cited.—Give only papers re-
ferred to in the text; list alphabetically by
author. Spell out references (journal names
etc.) completely, including conjunctions
and prepositions—no abbreviations. Do not
underline; journal names and book titles are
set in Roman type. See recent issues for
styles for journals, books, book chapters,
etc.
When referring to references in text, use
the following; Henry (1990), (Henry 1990),
(Henry 1987, 1990), (Smith 1990, Henry
1992); (South 1990; Henry 1990, 1991);
(Miller 1990a, 1990b) where two articles
are published by the same author in the
same year, or Henry et al. 1990 where more
592
than two authors are involved (do not ital-
icize ‘‘et al.’’). Citations ‘‘in press’’ should
be cited as ‘‘in press,’’ not with year of
expected publication.
Names of persons providing unpublished
information should include initials and be
referenced in the text as: W. Mathis (per-
sonal communication) [information ob-
tained orally] or W. Mathis (in litt.) [infor-
mation obtained in a writing].
Names of organisms.—The first men-
tion of a plant or animal should include the
full scientific name including the authority.
Use only common names approved in Com-
mon Names of Insects and Related Organ-
isms published by the Entomological Soci-
ety of America. Spell out the entire scien-
tific name the first time it is mentioned,
thereafter abbreviate generic names; how-
ever, do not abbreviate a genus name at the
beginning of a paragraph or sentence, or if
two or more genera being discussed have
the same first letter. Within sentences, al-
ways use the genus name or initial preced-
ing a species name.
Taxonomic papers.—Taxonomic papers
must conform to requirements of the Inter-
national Code of Zoological Nomenclature.
In addition, type specimens must be desig-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
nated for new species described, type de-
positories must be clearly indicated, and
new taxa must be clearly differentiated
from existing taxa by means of keys or dif-
ferential diagnoses. In short, these papers
must conform to good taxonomic practices.
Book Reviews.—Send to Book Review
Editor (see inside front cover) and see re-
cent Proceedings for format.
Charges.—Authors will be assessed a
charge to cover editing and publication ex-
penses. The current charge to members is
$40.00 per printed page. These charges are
in addition to those for reprints and author’s
correction in proof. The charge for pages in
excess of 15 printed pages, for non-mem-
bers, and papers for immediate publication
is $60.00 per printed page. Member authors
who are retired or not affiliated with an in-
stitution may request to have charges
waived and should do so when the manu-
script is submitted. A complete or partial
waiver may delay publication. Charges for
author errors and/or changes in proof, for
reprints if ordered, for immediate publica-
tion, and for non-members cannot be
waived. Acceptance of papers is based only
on scientific merit without regard to the au-
thor’s financial support.
PUBLICATIONS FOR SALE BY THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON
MISCELLANEOUS PUBLICATIONS
Cynipid Galls of the Eastern United States, by Lewis H. Weld. 124 pp. 1959 __
noid alls /of the Southwest, by Lewis H. Weld. 35 pp, 1960,
Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C. Steyskal.
SIS) RYO IGT IAT SS a ea ELE LEN
Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C.
SES SIT OIL fe) oY USI SB I SD a Na Eee
A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael
EL (SVCLMELUETE Ey betes liy 0) oS HS E210) eB is A CL Pa TP RS AE AG EEE ae ee ee
A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera): Second Edition, Revised, by
Erica Gnisselliand Michaelis chalittenc imp alloy yyersoe sek Tal CAT NUE ye Rota a al es le eae
MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
No.1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 __...
No. 2. A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving.
125): jo} Op A SY M0 peel BL Pol oe le Se ee
No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman.
APSNSI 0) oak LIAS £2 se a ace Ee GS Fd SR Ee SS SS BRR Th a SI A ee
No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952 ___-_-
No.5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 _..
No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi
Akahasiy 230) Pp. "LOGO x ete Le eae Ries | Neen ses eR ee Le ee RN Oe abe PV es
No. 7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp.
Ai ye Ly Ne G8 A AP TAG ED DJ DE IRN LE ee NEY Aiea De evn Ne oD a As MONEE INE ae Re
No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato-
pogonidae), by W. L. Grogan, Jr. and W.-W. Wirth. 125 pp. 1979
No.9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson.
PAULO): Foy OVE RS UA erat Ea I ae Poet oA ak RNS ene Eg | ee
No. 10. Recent Advances in Dipteran Systematics: Commemorative Volume in Honor of Curtis W.
Sabrosky. Edited by Wayne N. Mathis and F. Christian Thompson. 227 pp. 1982.
No. 11. A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370
PDE oe eee ere se me No Ae ee aE DV tk URS Wale es Ae gee a re ne ek A eI
No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff.
(Sy fo} ops ALS Loy IN a 2 Ae OO a A ee ORE LU Maan etl. hie ER armen ANON LS AOR ei 7 Sens 8
No. 13. An Identification Manual for the North American Genera of the Family Braconidae (Hyme-
noptera), by Paul M. Marsh, Scott R. Shaw, and Robert A. Wharton. 98 pp. 1987
No. 14. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174
Pp sel OO Shes area Bi DY aD TS NDE ae Oh Se Sed eine Ne Se ees Se ok Ee ae
No. 15. A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent
andeWe es Grogans UEslOS pp lOO spe tit cs os oe oN) Ee Ea ae ee eRe Le RL eee
No. 16. The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 __
No. 17. Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Krombein, edited
byebuis wNorden-andVAGS: Menke: 216) pps, 1996 je ae ae Ae ee
No. 18. Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and
Witham Gropamelte 297 spp lOO: ose es Eee a ae ee A en ee ae a rs te
25.00
Back issues of the Proceedings of the Entomological Society of Washington are available at $60.00 per volume
to non-members and $25.00 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, % Department of Entomology, MRC 168, Smithsonian
Institution, Washington, D.C. 20560.
CONTENTS
(Continued from front cover)
MOULTON, STEPHEN R., II and STEVEN C. HARRIS—New species of southwestern Ne-
arctic microcaddisflies (Trichoptera: Hydroptilidae)
RADOVIC, IVICA T. and SRDJAN SUSIC—Morphological characteristics of the sting and
prey carriage mechanism in Sericophorus relucens F. Smith (Hymenoptera: Sphecidae:
Larrinae)
ROBINSON, HAROLD and MARK DEYRUP—Two new species of Asyndetus Loew, and notes
on the identity of A. interruptus Loew (Diptera: Dolichopodidae)
SMITH, DAVID R. and TERENCE L. SCHIEFER—A new genus and species of Allantinae
(Hymenoptera: Tenthredinidae) from southeastern United States
SWEARINGEN, MICHEL, DAVID HEADRICK, and TOM BELLOWS—Comparison of fix-
ation and drying procedures for scanning electron microscopy among insect body types ...
TOGASHI, ICHIJI—A new genus and a new species belonging to the subfamily Blennocam-
pinae (Hymenoptera: Tenthredinidae) from Japan
WHEELER, A. G., JR. and E. RICHARD HOEBEKE—Trioza chenopodii Reuter: First North
American record of an Old World psyllid (Homoptera: Psylloidea: Triozidae)
WILKERSON, RICHARD C., MARIA ANICE M. SALLUM, and OSWALDO PAULO FOR-
ATTINI—Redescription of Anopheles (Anopheles) shannoni Davis; a member of the Arri-
balzagia series from the Amazon Basin (Diptera: Culicidae)
NOTES
EPSTEIN, MARC E.—Parasa indetermina (Biosduval) (Lepidoptera: Limacodidae), a new
host for Systropus macer Loew (Diptera: Bombyliidae)
MARSH, PAUL M.—Replacement names for Western Hemisphere genera of Doryctinae (Hy-
menoptera: Braconidae)
PAKALUK, JAMES and SANDRA L. ANAGNOSTAKIS—Acoptus suturalis LeConte (Co-
leoptera: Curculionidae: Zygopinae), a potential vector of the chestnut blight fungus, Cry-
phonectria parasitica (Murrill) Barr, in the eastern United States
PRICE, ROGER D. and RICARDO L. PALMA—Gender correction for a louse (Phthiraptera:
Menoponidae) from Cormorants
MISCELLANEOUS
ROBBINS, RICHARD G. and RALPH P. ECKERLIN—Robert Traub (1916-1996): An Ap-
preciation
INSTRUCTIONS FOR AUTHORS
a
i A te lg
* ,
ere ae A ee es
Ne eg
VOL. 99 OCTOBER 1997 NO. 4
(ISSN 0013-8797)
“! PROCEEDINGS
EN i of the
ENTOMOLOGICAL SOCIETY
of WASHINGTON
PUBLISHED
QUARTERLY
CONTENTS
BOHART, RICHARD M.—A review of the genus Hoplisoides Gribodo (Hymenoptera: Sphe-
Cidae Gry tit) MIT IN Orth FAMETLCAN AUN Ly, Mires. co sigs crcseia te sia cise Gatercle peels ste sacle ew sieie wean eteaaete 645
BYERS, GEORGE W.—Four puzzling new species of Mecoptera ............5......2e eee eeee neces 681
CONDON, MARTY, ANN TRUELOVE, and LYNN MATHEW—Sexual dimorphism in
mouthparts of Blepharoneura Loew (Diptera: Tephritidae) ....................... 0... s cece eee 676
GOEDEN, RICHARD D. and JEFFREY A. TEERINK—Life history and description of imma-
ture stages of Xenochaeta dichromata Snow (Diptera: Tephritidae) on Hieracium albiflorum
Hooker inecentral and:southern Calitormiay.(0\2 Cie eos iose toe hate pate Rcscipe oy tanaeletictecee oa epee alae 597
GOEDEN, RICHARD D. and JEFFREY A. TEERINK—Life history and description of imma-
ture stages of Trupanea signata Foote (Diptera: Tephritidae) on Gnaphalium luteo-album
IesN SOUTMSrMUC ab fORMMAy. a. skis ss sors see AR tee tah Reta te eae es od ne ans cls biel see casos aac tn eer eams 748
GRISSELL, E. E.—Biological notes on Sparasion Latreille (Hymenoptera: Scelionidae), an egg
parasitoid of Atlanticus gibbosus Scudder (Orthoptera: Tettigonidae) .......................... 693
JONES, ROBERT W. and HORACE R. BURKE—New species and host plants of the Antho-
nomus grandis species group (Coleoptera: Curculionidae) ............... 2.2222 ee cece e rece eens 705
KOTRBA, MARION—Shoot or stab? Morphological evidence on the unresolved oviposition
technique in Stylogaster Macquart (Diptera: Conopidae), including discussion of behavioral
OHSELVAGLOMS year AROS Mis eae sa ahaiz chert Situs cles NAD aaaG Sree TEETER o eee OU Mee EL RE a RTL LLes fe aiabe a 614
MARTINEZ M., IMELDA and VICENTE HERNANDEZ-ORTIZ—Anatomy of the reproduc-
tive system in six Anastrepha species and comments regarding their terminology in Te-
Pumiidacs(Diptera) ie ste Mews Alne ucla nese. aah dee aM ern tee See OR i REE A OM BAL NEP fe 724
MARSHALL, S. A. and S. FITZGERALD—Rudolfina cavernicola, a new species of cave-
associated Sphaeroceridae (Diptera) from Colorado and Arizona ..................2.22.0022005- 641
MATHIS, WAYNE N.—A revision of Neotropical Ditrichophora Cresson (Diptera: Ephyd-
LPI P=) Mio penta el te al nr Bib A Oa eI RU ICA Ar old airy eA Pa aba Ve ee Og 697
(Continued on back cover)
THE
ENTOMOLOGICAL SOCIETY
OF WASHINGTON
ORGANIZED MARCH 12, 1884
OFFICERS FOR 1997
M. ALMA SOLIs, President MICHAEL G. POGUE, Treasurer
WARREN E. STEINER, JR., President-Elect Davip G. FuRTH, Program Chair
HAROLD HARLAN, Recording Secretary JOHN W. Brown, Membership Chair
Ho.Luis B. WILLIAMS, Corresponding Secretary RALPH P. ECKERLIN, Past President
GarRY L. MILLER, Custodian
DAVID R. SmiTH, Editor
Publications Committee
THOMAS J. HENRY WAYNE N. MATHIS
Gary L. MILLER, Book Review Editor
Honorary President
CurTis W. SABROSKY
Honorary Members
LoulIsE M. RUSSELL ALAN STONE KARL V. KROMBEIN
All correspondence concerning Society business should be mailed to the appropriate officer at the following
address: Entomological Society of Washington, % Department of Entomology, MRC-168, Smithsonian Insti-
tution, Washington, D.C. 20560.
MEETINGS.—Regular meetings of the Society are held in the Natural History Building, Smithsonian Institu-
tion, on the first Thursday of each month from October to June, inclusive, at 8 P.M. Minutes of meetings are
published regularly in the Proceedings.
MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology.
Annual dues for members are $25.00 (U.S. currency).
PROCEEDINGS.—The Proceedings of the Entomological Society of Washington (ISSN 0013-8797) are pub-
lished quarterly beginning in January by The Entomological Society of Washington. POSTMASTER: Send
address changes to the Entomological Society of Washington, % Department of Entomology, MRC-168, Smith-
sonian Institution, Washington, D.C. 20560. Members in good standing receive the Proceedings of the Ento-
mological Society of Washington. Nonmember U.S. subscriptions are $60.00 per year and foreign subscriptions
are $70.00 per year, payable (U.S. currency) in advance. Foreign delivery cannot be guaranteed. All remittances
should be made payable to The Entomological Society of Washington.
The Society does not exchange its publications for those of other societies.
PLEASE SEE PP. 615-616 OF THE JULY 1996 ISSUE FOR INFORMATION 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, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash-
ington, D.C. 20560.
Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology,
Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560.
Books for Review: Gary L. Miller, Systematic Entomology Laboratory, ARS, USDA, Building 046, BARC-
West, Beltsville, MD 20705.
Managing Editor and Known Bondholders or other Security Holders: none.
This issue was mailed 24 October 1997
Second Class Postage Paid at Washington, D.C. and additional mailing office.
PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA
This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 593-596
TRICHOGRAMMATOIDEA BRASILIENSIS (ASHMEAD)—NEW
COMBINATION FOR A SPECIES HISTORICALLY PLACED IN
TRICHOGRAMMA (HYMENOPTERA: TRICHOGRAMMATIDAE)
JOHN D. PINTO
Department of Entomology, University of California, Riverside, CA 92521, U.S.A.
Abstract.—The species brasiliensis Ashmead is transferred from Trichogramma to
Trichogrammatoidea. This new combination is based on the examination of the re-
mounted holotype.
Key Words.
In 1904 Ashmead described the species
Pentarthron brasiliensis from a single fe-
male “‘collected on cotton” in “‘Bahia’’,
Brazil, in ‘““Mar 83’’. Pentarthron Riley has
long been considered a synonym of Trich-
ogramma Westwood, and brasiliense ' has
always been associated with these nominal
genera (Zerova and Fursov 1989). Because
the identification of Trichogramma species
depends largely on male traits, it was point-
ed out by Pinto and Stouthamer (1994) that
the definition of Trichogramma brasiliense
was unresolved, notwithstanding its contin-
ued use in the biological control literature
(see De Santis 1989 and below for refer-
ences). Upon examining the female holo-
type in the National Museum of Natural
History, Smithsonian Institution (Washing-
ton, D.C.) (USNM type no. 6596) the ques-
tion arose as to whether brasiliense was
correctly placed in Trichogramma. Unfor-
tunately, the type was uncleared and poorly
mounted, precluding examination of certain
fore- and hind wing characters critical for
positive generic placement. The specimen
' Because this specific name has been associated
with generic names of different gender (Penarthron,
Trichogramma, and Trichogrammatoidea), its ending
varies in this paper depending on the one it is associ-
ated with in discussion.
Trichogramma, Trichogrammatoidea, taxonomy
was recently remounted and it is now clear
that the species is not a Trichogramma but
instead belongs to the related genus Tricho-
grammatoidea Girault.
Trichogrammatoidea is most easily sep-
arated from Trichogramma by male geni-
talia and antennae (Pinto and Stouthamer
1994). However, certain wing features al-
low either sex to be placed. Trichogramma
(Fig. 1) is characterized as follows: Fore-
wing (Fig. 1a) marginal vein with three ro-
bust and elongate setae on dorsal surface;
premarginal vein with two such setae. Two
suboval sensilla between the marginal and
premarginal veins. RS, vein track present
behind stigmal vein. Hind wing (Fig. 1b)
usually with at least two setal tracks, a mid-
dle track which is always complete to the
wing apex and a complete or partial pos-
terior track; a third, anterior track is present
or absent. In contrast, Trichogrammatoidea
is characterized as follows: Forewing mar-
ginal vein with only two robust and elon-
gate setae on the dorsal surface—the third
(apical most) is considerably shorter; pre-
marginal vein with only one seta. One sub-
oval sensillum between marginal and pre-
marginal veins. RS, vein track absent. Hind
wing with only a middle setal track which
is not complete to the wing apex. In addi-
594
la.
2a.
025mm. b. s&
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-2. Wing characteristics. 1, Trichogramma: a, Forewing venation, dorsal; b, hind wing, anterior at top.
2, Holotype female of Trichogrammatoidea brasiliensis: a, forewing venation, dorsal; b, hind wing, anterior at
top. Abbreviations: m = marginal vein; p = premarginal vein; arrow refers to boundary between marginal and
premarginal veins.
tion, the forewing is generally broader in
Trichogramma than in Trichogrammato-
idea and it has denser setation and shorter
fringe setae along its margin.
The holotype of brasiliensis is clearly as-
signable to Trichogrammatoidea based on
wing characters. During remounting, one
forewing and hind wing were placed to-
gether under a separate coverslip for opti-
mal viewing and illustration (Fig. 2a, b). In
this specimen the RS, vein track is absent;
only a single setal socket occurs on the pre-
marginal vein, and of the three setae on the
marginal vein, the two basal ones are bro-
ken but the third, which is present, is rela-
tively short as in Trichogrammatoidea. The
single suboval sensillum between the pre-
marginal and marginal veins is not visible
in the remounted forewing of the type, but
it can be seen on the wing which remains
attached to the body. Also, as is character-
istic of Trichogrammatoidea but not Trich-
ogramma there is only a single, incomplete
setal track on the hind wing of the type.
Although a single track occurs in two spe-
cies of the primitive Lachesis Group of
Trichogramma (Pinto 1992), it is complete
to the wing apex in all species. In addition,
the forewing of the type has relatively
sparse setation and an elongate setal fringe
VOLUME 99, NUMBER 4
on its margin, both characteristic of Tricho-
grammatoidea. It is curious that Girault
(1911b) upon redescribing this species did
not recognize its incorrect generic place-
ment considering that he described Tricho-
grammatoidea earlier in the same year (Gi-
rault 191la) and noted the unique wing
characteristics in his redescription (absence
of an RS, and single setal track in hind
wing) which he had incorporated into the
definition of the new genus.
Because the holotype of Trichogramma-
toidea brasiliensis (new combination) is a
female and in poor condition I am unable
to determine with confidence if it is distinct
or a synonym of an already described spe-
cies. It is possible that Trichogrammatoidea
annulata De Santis, also associated with
cotton in Brazil (De Santis 1972), is a syn-
onym of T. brasiliensis. | have examined a
paratype male and female of 7. annulata
and can find no differences in wing struc-
ture or Ovipositor length. However, until the
South American Trichogrammatoidea are
better known I suggest continuing to use T.
annulata as a valid name and treating T.
brasiliensis as valid but as a nomen dubium,
i.e. a name not certainly applicable to any
known taxon.
The question arises as to what species of
Trichogramma has/have been misidentified
as brasiliense in the literature where the
name continues to be associated with bio-
logical control efforts particularly against
pyralid pests of sugarcane (Monje 1995).
This is not possible to answer without ex-
amining voucher specimens associated with
each study. However several years ago I re-
ceived two cultures identified as Tricho-
gramma brasiliense, one from the USDA
Laboratory in Beltsville, Maryland, and the
other from a Trichogramma_ production
center in Torreon, Mexico. Both were T.
fuentesi Torre, a species generally misiden-
tified as T. fasciatum Perkins in the early
literature. Trichogramma fuentesi was orig-
inally described from Cuba but it also oc-
curs in the southeastern United States, Mex-
ico and South America (Pinto et al., 1983).
595
The literature suggests that the name bra-
siliense has been applied to other species as
well. For example, Ruiz and Korytkowski
(1980) define it as a species which I would
consider close to 7. pretiosum based on
their description and illustrations, and Kfir
(1982) and others have applied the name to
thelytokous populations. The latter usage
may stem from Quednau (1961), and Na-
garaja and Nagarkatti (1969) who consid-
ered brasiliense as a thelytokous form of
Trichogramma fasciatum. The actual spe-
cies studied by Kfir, Ruiz and Korytkowski,
and several other authors, remain undeter-
mined. One of the mysteries of Tricho-
gramma taxonomy is how certain common
New World species became misidentified as
they did and how these misidentifications
became so well established in the literature.
ACKNOWLEDGMENTS
The type of Trichogrammatoidea bra-
siliensis was remounted by Gary Platner.
Figures 1 and 2 were prepared by Marina
Planoutene. I thank Andrew Polaszek for
advice on remounting procedures, and Mi-
chael Schauff (Systematic Entomology
Laboratory, USDA) for the loan of the type
and granting permission to remount it.
LITERATURE CITED
Ashmead, W. H. 1904. Classification of the chalcid
flies. Memoirs of the Carnegie Museum Vol. | (4),
Sepp:
De Santis, L. 1972. Un nuevo tricogrammatido
(Hym.) neotropico parasito de los huevos de Al-
abama agrillacea (Lep.). Arquivos do Instituto
Bioldgico, Sao Paulo 39: 121-124.
De Santis, L. 1989. Catalogo de los himenopteros cal-
cidoideos (Hymenoptera) al sur de los estados uni-
dos. Segundo Suplimento. Acta Entomologica
Chilena 15: 9-90.
Girault, A. 191la. Descriptions of nine new genera of
the chalcidoid family Trichogrammatidae. Trans-
actions of the American Entomological Society
37: 1-41, 1 pl.
Girault, A. 1911b. Synonymic and descriptive notes
on the chalcidoid family Trichogrammatidae with
descriptions of new species. Transactions of the
American Entomological Society 37: 43-83.
Kfir, R. 1982. Reproduction characteristics of Trich-
ogramma brasiliensis and T. lutea, parasitising
596
eggs of Heliothis armiger. Entomologia Experi-
mentalis et Applicata 32: 249-255.
Monje, J. C. 1995. Present significance of Tricho-
gramma spp. (Hymenoptera: Trichogrammatidae)
for the control of sugarcane borers in the Ameri-
cas. Mitteilungen der Deutschen Gesellschaft fiir
allgemeine und angewandte Entomologie 10:287—
290.
Nagaraja, H. and S. Nagarkatti. 1969. Three new spe-
cies of Trichogramma [Hymenoptera: Tricho-
grammatidae] from India. Entomophaga 14: 393—
400.
Pinto, J. D. 1992. Novel taxa of Trichogramma from
the New World tropics and Australia (Hymenop-
tera: Trichogrammatidae). Journal of the New
York Entomological Society 100: 621—633.
Pinto, J. D. and R. Stouthamer. 1994. Systematics of
the Trichogrammatidae with emphasis on Tricho-
gramma. Chap. 1, pp. 1-36. In, Wajnberg, E. and
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
S. A. Hassan, eds. Biological control with egg par-
asitoids. CAB International, Wallingford.
Pinto, J. D., E. R. Oatman and G. R. Platner. 1983.
The identity of two closely related and frequently
encountered species of New World Trichogramma
(Hymenoptera: Trichogrammatidae). Proceedings
of the Entomological Society of Washington 85:
588-593.
Quednau, W. 1961. Die Problematik der Nomenklatur
bei den Trichogramma—Arten. Entomophaga 6:
155-161.
Ruiz, E. R. and C. A. Korytkowski. 1979. Contribu-
cion al conocimiento de los Trichogrammatidae
(Hymenoptera: Chalcidoidea) del Peru. Revista
Peruana de Entomologia 22: 1-8.
Zerova, M. D. and V. N. Fursov. 1989. A catalogue
of species of the genus Trichogramma Westwood
of the world (Hymenoptera, Trichogrammatidae).
Institute of Zoology of the Science Academy of
the Ukrainian SSR. Publication 89 (4), 52 pp.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 597-607
LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF
XENOCHAETA DICHROMATA SNOW (DIPTERA: TEPHRITIDAE) ON
HIERACIUM ALBIFLORUM HOOKER IN CENTRAL AND
SOUTHERN CALIFORNIA
RICHARD D. GOEDEN AND JEFFREY A. TEERINK
Department of Entomology, University of California, Riverside, CA 92521, U.S.A.
Abstract.—Xenochaeta dichromata Snow is a nearly monophagous, uni- or bivoltine,
synovigenic, seed-feeding fruit fly attacking the flower heads of certain native hawkweeds
(Hieracium spp., Asteraceae) in western North America. Xenochaeta aurantiaca (Doane)
is synonymized with X. dichromata. The egg, first-third instar larvae, and puparium are
described and all but the second instar are illustrated. The third instar is shagreened, i.e.,
meso-, metathorax, and abdominal segments are covered by minute, dome-shaped ver-
rucae. Xenochaeta dichromata differs from other shagreened species in possessing a single
row of verruciform sensilla circumscribing each thoracic and abdominal segment. The life
cycle is of the aggregative type. Eggs are deposited singly in individual immature flower
heads. Each larva consumes the entire contents of a single head, within which it over-
winters as a prepuparium. Pupariation occurs in the spring and the newly emerged adults
aggregate on the preblossom host plants to mate and oviposit. The possible use of this
fly for biological control of hawkweeds in North America and abroad, e.g., in New Zea-
land, is discussed.
Key Words: Insecta, Xenochaeta, Hieracium, hawkweeds, biology, taxonomy of im-
mature stages, florivory, monophagy, reproductive behavior, parasitoids, bi-
ological control of weeds
The rearing of a good series of females
that key to Xenochaeta aurantiaca (Doane)
flower head samples collected in June or
August from different locations on the
and males that key to X. dichromata Snow
from the same samples of mature flower
heads of Hieracium albiflorum Hooker (As-
teraceae) in southern California confirm
that there is a single, sexually dimorphic
species of Xenochaeta. These results pro-
vided us the opportunity to synonymize
these tephritid species and to study the life
history and describe the immature stages of
this heretofore-little-known, nonfrugivorous
fruit fly.
MATERIALS AND METHODS
Our field studies on X. dichromata fo-
cused mainly on laboratory dissections of
western slopes of the Sierra Nevada Moun-
tains in central and southern California dur-
ing 1993-1995: (1) N of Union Valley Res-
ervoir at 1700-m elevation, Eldorado Na-
tional Forest, Eldorado Co., 19.1x.1993; (2)
four sites between 1820 and 2050 m in
Mountain Home State Forest, Tulare Co.,
3—4.1x.1994; (3) S of Slate Mountain and E
of Onion Meadow at 2120 m, Sequoia Nat.
Forest (N-section), Tulare Co., 3.ix.1994;
(4) S of Sampson Flat at 1210 m, Sequoia
Nat. Forest (N-section), Fresno Co.,
14.vi.1995. These locations were too far
from Riverside to allow field observation of
598
adult behaviors. Samples of immature or
mature flower heads of H. albiflorum were
transported in cold-chests in an air-condi-
tioned vehicle to the laboratory and stored
under refrigeration for subsequent dissec-
tion, photography, description, and mea-
surement. Thirty eggs, five first-, 12 sec-
ond-, and 18 third-instar larvae, and six pu-
paria dissected from flower heads were pre-
served in 70% EtOH for scanning electron
microscopy (SEM). Additional mature lar-
vae and prepuparia were placed in separate,
glass shell vials stoppered with absorbant
cotton and held in humidity chambers at
room temperature for adult and parasitoid
emergence. In 1994, those larvae and pre-
puparia not pupariating by mid-December
were stored in a refrigerator at 2 + 1°C until
August, 1995, before returning them to hu-
midity chambers. Specimens for SEM were
hydrated to distilled water in a decreasing
series of acidulated EtOH. They were os-
micated for 24 h, dehydrated through an in-
creasing series of acidulated EtOH and two,
1-h immersions in Hexamethlydisilazane
(HMDS), mounted on stubs, sputter-coated
with a gold-palladium alloy, and studied
with a JEOL JSM C-35 SEM in the De-
partment of Nematology, University of Cal-
ifornia, Riverside.
Most adults reared from isolated puparia
were individually caged in 850-ml, clear-
plastic, screened-top cages with a cotton
wick and basal water reservoir and provi-
sioned with a strip of paper toweling im-
pregnated with yeast hydrolyzate and su-
crose. These cagings were used for longev-
ity studies in the insectary of the Depart-
ment of Entomology, University of
California, Riverside, at 25 + 1°C, and
14/10 (L/D) photoperiod. Virgin male and
female flies obtained from emergence vials
were paired in clear-plastic petri dishes pro-
visioned with a flattened, water-moistened
pad of absorbant cotton spotted with honey
(Headrick and Goeden 1991, 1994) for di-
rect observations, videorecording, and still-
photography of their courtship and copula-
tion behavior.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Plant names used in this paper follow
Munz (1974); tephritid names and adult ter-
minology follow Foote et al. (1993). Ter-
minology and telegraphic format used to
describe the immature stages follow Goe-
den et al. (1993, 1994a, b, 1995a, b) and
Goeden and Teerink (1996a, b, c; 1997) and
our earlier works cited therein. Means + SE
are used throughout this paper. Voucher
specimens of X. dichromata and its parasit-
oids reside in the research collections of
RDG; preserved specimens of eggs, larvae
and puparia are stored in a separate collec-
tion of immature Tephritidae maintained by
JAT.
RESULTS AND DISCUSSION
TAXONOMY
Adult.—Xenochaeta dichromata and X.
aurantiaca (Doane) were termed “rarely
collected’”’ tephritids by Foote et al. (1993).
Xenochaeta dichromata was described from
a single male (therefore holotype) by Snow
(1894), and besides records for additional
males, only a single female specimen has
since been recorded (Foote and Blanc 1979,
Foote et al. 1993). Xenochaeta aurantiaca
was described (as a Eutreta) by Doane
(1899) and only females heretofore were
known. Noting these disparities in sexes re-
corded, Foote et al. (1993) suggested that
‘*.. further studies may show these two
species to be conspecific.”” As predicted,
only one, sexually dimorphic species is
present in California, and these two names
are synonyms. Both males of X. dichromata
(n = 34) and females that keyed to X. au-
rantiaca (n = 29) were consistently and
solely reared together from single samples
of mature flower heads of H. albiflorum
collected in August at the above-named
study sites, and shortly after their puparia-
tion and emergence as adults, these flies
readily mated when placed together in petri
dish arenas (n = 5, see below). Therefore,
X. aurantiaca is hereby synonymized with
X. dichromata, which has priority.
The wing patterns of both sexes of X.
VOLUME 99, NUMBER 4
Riss 1
Egg of X. dichromata. pedicel to left.
dichromata were pictured in Foote et al.
(1993), the female as aurantiaca, and the
male wing pattern was figured in Foote and
Blanc (1963).
Immature stages.—Egg.: Twenty eggs of
X. dichromata were white, opaque, smooth;
with an elongate-ellipsoidal body, 0.44 +
0.002 (range, 0.41—0.46) mm long, 0.17 +
0.002 (range, 0.17—0.19) mm wide, smooth-
ly rounded at tapered posterior end, and
with a peg-like anterior pedicel, 0.02 mm
long (Fig. 1).
Third instar: Oblong-ovoid, tapering an-
teriorly, truncated posteriorly, shagreened
(Fig. 2A); gnathocephalon conical, smooth,
few rugose pads laterally (Fig. 2B); dorsal
sensory organ a single, dome-shaped papil-
la (Fig. 2B-1, C-1); anterior sensory lobe
(Fig. 2B-2, C) bears terminal sensory organ
(Fig. 2C-2), pit sensory organ (Fig. 2C-3),
lateral sensory organ (Fig. 2C-4) and su-
pralateral sensory organ (Fig. 2C-5); stomal
sense organ, a distinct lobe invested with
sensoria ventrolaterad of anterior sensory
lobe (Fig. 2B-3, D-1); mouth hooks triden-
tate (Fig. 2B-4, D-2); median oral lobe ta-
pers anteriorly, attached to labial lobe (Fig.
2D-3); prothorax circumscribed anteriorly
with minute acanthae, single row of verru-
ciform sensilla circumscribe each thoracic
and abdominal segment; anterior thoracic
spiracles with 4—6 raised papillae (Fig. 2E);
meso- and metathoracic lateral spiracular
complexes consist of an open spiracle and
599
a single verruciform sensillum; abdominal
lateral spiracular complexes consist of an
open spiracle (Fig. 2F-1) and two verruci-
form sensilla (Fig. 2F-2); posterior spirac-
ular plates consist of three ovoid rimae, ca.
0.03 mm in length (Fig. 2G-1), and four
interspiracular processes, longest measuring
0.01 mm (Fig. 2G-2); verruciform sensilla
circumscribe caudal segment (Fig. 2G-3);
compound sensilla each consist of two ver-
ruciform sensilla, one with a central papilla
(Fig. 2G-4, H).
Norrbom et al. (1997) placed Xenochaeta
in the new Tribe Noeetini along with Aci-
dogona and Jamesomyia. Benjamin (1934)
briefly described the immature stages of
Acidogona melanura Loew, and the general
habitus of the egg, mature larva, and pu-
parium of this species do resemble X. di-
chromata. However, X. dichromata is the
only species in the tribe for which the im-
mature stages are described in detail.
Xenochaeta was placed in the Tribe Eu-
tretini by Foote et al. (1993). Other genera
and species in this tribe for which the im-
mature stages have been described in sim-
ilar detail include Eutreta diana (Osten
Sacken) (Steck and Wharton 1986; Goeden
1990a, b) and Paracantha gentilis Hering
(Headrick and Goeden 1990a). Xenochaeta
dichromata differs from these two species
in having a shagreened mature larva. Goe-
den (1990a) described E. diana (Osten
Sacken) first instar as ringed with verrucae
on abdominal segments II—V. However, the
verrucae are absent or nearly so on the sec-
ond and third instars (Steck and Wharton
1986, Goeden 1990a). Two species from
other tribes in which the mature larva also
are shagreened are Tomoplagia cressoni
Aczel (Tribe Acrotaeniini) and Valentibulla
californica (Coquillett) (Tribe Dithrycini)
(Goeden and Headrick 1991, Goeden et al.
1995b, Foote et al. 1993). The latter of
these two species most closely resembles X.
dichromata, but lacks the verruciform sen-
silla that circumscribe the thoracic and ab-
dominal segments in X. dichromata. Also,
the posterior two-thirds of the prothorax is
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Third instar of X. dichromata. (A) habitus, anterior to left; (B) gnathocephalon, anterior view, 1—
dorsal sensory organ, 2—anterior sensory lobe, 3—stomal sense organ, 4—mouth hooks; (C) anterior sensory
lobe, 1—dorsal sensory organ, 2—terminal sensory organ, 3—pit sensory organ, 4—lateral sensory organ, 5—
supralateral sensory organ; (D) gnathocephalon lateral view, 1—stomal sense organ, 2—mouth hooks, 3—median
oral lobe; (E) anterior thoracic spiracle; (F) fourth abdominal lateral spiracular complex, 1—spiracle, 2—ver-
VOLUME 99, NUMBER 4
lene, Sie
anterior to left; (B) gnathocephalon, anterior view, 1—
dorsal sensory organ, 2—anterior sensory lobe, 3—
mouth hooks; (C) caudal segment, posterior spiracular
plates.
First instar of X. dichromata. (A) habitus,
ae
601
shagreened in V. californica, the mouth
hooks are bidentate, and the abdominal lat-
eral spiracular complex consists of an open
spiracle and a single sensillum (Goeden et
al. 1995b).
Second instar: Elongate-ovoidal, round-
ed anteriorly and posteriorly, shagreened;
gnathocephalon conical, few rugose pads
laterad of stomal sense organ; dorsal sen-
sory organ composed of a single papilla;
anterior sensory lobe bears all four sensory
organs; stomal sense organs distinct, ven-
trolaterad of anterior sensory lobe; mouth
hooks bidentate; median oral lobe tapering
anteriorly; thoracic segments smooth, single
row of verruciform sensilla circumscribe
thoracic and abdominal segments; anterior
thoracic spiracles and lateral spiracular
complex were not observed; posterior spi-
racular plates bear three ovoid rimae, Ca.
0.018 mm in length and four interspiracular
processes, longest measuring 0.006 mm;
compound sensilla were not observed.
The second instar differs from the mature
larva in that the dome-shape verrucae are
smaller and restricted to the dorsad and in-
tersegmental lines of the abdominal seg-
ments, and the mouth hooks are bidentate.
First instar: Elongate-ellipsoidal, minute
verrucae circumscribing abdominal seg-
ments (Fig. 3A); gnathocephalon conical,
smooth, lacking rugose pads (Fig. 3B); dor-
sal sensory organ consists of a dome-
shaped papilla (Fig. 3B-1); anterior sensory
lobe bears all four sensory organs (Fig.
3B-2); stomal sense organs indistinct;
mouth hooks bidentate (Fig. 3B-3); median
oral lobe not observed; prothorax with mi-
nute acanthae ventrally; posterior spiracular
plates contiguous, bear two ovoid rimae and
four interspiracular processes (Fig. 3C).
The first instar has fewer, smaller, dome-
shaped verrucae than later instars. The first
instar habitus is more ellipsoidal in shape,
ruciform sensilla; (G) caudal segment, |—rima, 2—interspiracular process, 3—verruciform sensillum, 4—com-
pound sensillum; (H) posterior compound sensillum, verruciform sensillum with central papilla.
602
not as tapered anteriorly. Sensory structures
such as the stomal sense organ are not as
well-defined in the first instar as they are in
the later instars. The verruciform sensilla
circumscribing the thoracic and abdominal
segments in the later instars were not ob-
served.
Puparium: Dark brown, elongate-ellip-
soidal, tapering anteriorly, rounded poste-
riorly, shagreened (Fig. 4A); anterior end
bears invagination scar (Fig. 4B-1) and an-
terior thoracic spiracles (Fig. 4B-2); caudal
segment bears posterior spiracular plates
(Fig. 4C-1), compound sensilla (Fig. 4C-2),
and verruciform sensilla (Fig. 4C-3). Twen-
ty-five puparia of X. dichromata averaged
3.19 + 0.05 (range, 2.85—3.60) mm in
length; 1.61 + 0.03 (range, 1.48—1.79) mm
in width.
DISTRIBUTION AND Hosts
The distribution of X. dichromata in
North America north of Mexico was
mapped by Foote et al. (1993) (also as X.
aurantiaca) to include locations in British
Columbia, California, Montana, Oregon,
Utah, and Washington. Linda Wilson and J.
McCaffrey (in litt. 1995, 1996) also have
reared X. dichromata from Hieracium al-
biflorum and the H. scouleri Hooker com-
plex in Idaho, but not from any of the in-
troduced hawkweeds, e.g., H. pratense
Tausch. Between 1987 and 1994, eight,
l1-liter samples of mature flower heads of
H. albiflorum collected by RDG (unpub-
lished data) at locations in El] Dorado, Fres-
no, Kern, Madera, Mono, and Trinity coun-
ties different from our above-named study
sites all failed to yield X. dichromata. Sim-
ilarly, six samples of mature flower heads
of H. horridum Fries collected by RDG
(unpublished data) from 1987-1993 yielded
no X. dichromata. Only one other host-
plant has been reported, H. cynoglossoides
Arvet-Touvet (Novak et al. 1967, Foote et
al. 1993), indicating that X. dichromata is
nearly monophagous (one host genus) in
flower heads of certain species of Hiera-
cium, but does not occur in all parts of its
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 4.
Puparium of X. dichromata. (A) habitus,
anterior to right; (B) anterior end, 1—invagination
scar, 2—anterior thoracic spiracle; (C) caudal segment,
1—posterior spiracular plates, 2—compound sensilla,
3—verruciform sensillum.
host-plants’ geographic ranges. Dodson and
George (1986) demonstrated convincingly
that the gall-forming tephritid flies Aciurina
bigeloviae (Cockerell) and A. trixa Curran
VOLUME 99, NUMBER 4
are less widely distributed than their host
plants within four southwestern States.
BIOLOGY
Egg.—Only two (6%) of 34 preblossom
flower heads contained two eggs each, the
rest of these closed, immature heads each
contained a single egg (Fig. 5A, B). Im-
mature heads with eggs averaged 1.07 +
0.04 (range, 0.64—1.50) mm in diameter.
Eleven (32%) of 34 eggs examined were
oriented with the long axis parallel to the
long axes of the tiny, immature florets (Fig.
5A), 14 eggs (41%) were inserted nearly
perpendicular to the long-axes (Fig. 5B)
and nine eggs (26%) were inserted at angles
of 45—60°. Only one (9%) of the 11 eggs
inserted parallel to the florets (pedicel-last
like all tephritid eggs examined by us to
date, Fig. 5A), penetrated an ovule or floral
tube; whereas 14 (60%) of the 23 eggs ovi-
posited at an angle to the florets penetrated
an ovule or floral tube after the aculeus had
penetrated one or more phyllaries during
oviposition. Ten eggs rested atop the im-
mature florets and parallel to the receptacle
in the small cavity formed by the overlap-
ping phyllaries (Fig. 5B).
Larva.—Newly eclosed first instars (Fig.
5C) fed parallel to the receptacle on ovules
or floral tubes inside closed heads that av-
eraged 1.36 = 0.08 (range, 1.14—1.71; n =
7) mm in diameter. Six first instars each
damaged an average of 2.5 + 0.7 (range,
1—5) florets. The second instar (Fig. 5D) fed
centrally, well above and parallel to the re-
ceptacle, consuming the upper parts of the
ovules and the bases of the floral tubes
within a circle of intact, outer florets (Fig.
5D). Eleven feeding chambers of second in-
stars within closed, immature, flower heads
averaging 2.47 + 0.14 (range, 1.71—3.13)
mm in diameter measured 1.32 + 0.12
(range; 0:85—1:71)) mm by 1.03. = 0.13
(range, 0.57—1.42) mm (Fig. 5D).
The third instar continued to enlarge the
feeding chamber, eventually destroying all
ovules and floral tubes within the infested
heads (Fig. 5E, F). Thirty uninfested mature
603
heads of H. albiflorum produced an average
of 19 + 0.6 (range, 15—28) achenes. Usu-
ally, only one larva developed to maturity
in each of 132 infested heads examined
(Fig. 5E). Once the third instar consumed
the florets, it deeply scored the receptacle
and supplemented its diet with sap that col-
lected in the feeding depression, as reported
with several, but not all, florivorous, Cali-
fornia Tephritidae that we have studied
(Goeden 1988, Headrick and Goeden
1990a, b, Goeden and Headrick 1991, 1992,
Goeden et al. 1993, 1995a, Headrick et al.
1996). The infested heads never opened or
showed any protruding pappus apically and
were more conical in shape than the ellip-
soidal, uninfested heads (Fig. 5F). One hun-
dred heads containing unparasitized mature
larvae or prepuparia averaged 3.26 + 0.05
(range, 2.00—4.57) mm in diameter (Fig.
5F). The feeding cavities of these fully
grown, third instars measured 3.91 + 0.05
(range, 1.71—5.13) mm in length by 2.23 +
0.03 (range, 1.42—2.85) mm in width (Fig.
5E). The walls of these chambers were stiff-
ened by a dried, hardened mixture of floret
fragments, feces, and dried sap that coated
and glued together the inner walls of the
phyllaries (Fig. 5E).
Pupa.—Fully grown larvae ceased feed-
ing, contracted their bodies, invaginated
their mouthparts, and darkened ventrally
and posteriorly to become prepuparia. Most
prepuparia entered diapause and overwin-
tered in the dried heads, but eight of 33
(25%) held in moist chambers in the labo-
ratory in 1994 reversed their position 180°
within excised flower heads, pupariated
(Fig. 5G), and emerged as adults (Fig. 5H,
I).
Adult.—Foote and Blanc (1979) re-
marked that the entire genus Xenochaeta at
that time was represented in U.S. collec-
tions by a total of only 10—12 specimens.
After 3 years of study, our reared specimens
from California number 63, of which 34
(54%) are males. Adults were long-lived
under insectary conditions, as 13 males av-
eraged 48 + 7 (range, 15-97) days, and 12
604 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 5. Life stages of X. dichromata on Hieracium albiflorum. (A) egg (arrow) inserted between florets of
immature, preblossom flower head; (B) egg (arrow) resting atop florets parallel to receptacle; (C) first instar
feeding at base of floral tube; (D) second instar feeding on ovules and floral tube bases; (E) fully grown third
instar in feeding chamber; (F) infested flower head containing third instar (left), uninfested head right; (G)
Puparium in head; (H) mating pair, lateral view; (I) mating pair, ventral view. Lines = 1 mm.
VOLUME 99, NUMBER 4
females averaged 31 + 5 (range, 8—65)
days. Males (n = 5) readily mated when 3
to 18 days old; females (n = 5), when 3 to
17 days old (Fig. 5H, I). A 20-day-old fe-
male contained a full compliment of full-
size ova, but otherwise ovigenesis and ovi-
position were not studied in the laboratory
or field, nor were free-living adults ever ob-
served or collected.
As observed with adults of Eutreta diana
by Goeden (1990a, unpublished data) and
Paracantha gentilis (Headrick and Goeden
1990a, 1995), adults characteristically rest-
ed or walked with their wings arched and
parted and exhibited abdominal flexures.
Another behavior exhibited by males of X.
dichromata and E. angusta Banks (Head-
rick and Goeden 1995) was side-stepping
displays by males when facing and tracking
females in arenas. Wing displays by males
and females of X. dichromata most com-
monly were synchronous supinations, and
less commonly, slower paced, asynchro-
nous wing supinations by males, or wing
lofting of 20° embellished with rapid wing
vibration, when tracking females. No ab-
dominal pleural expansion by males was
observed, and also unlike P. gentilis (Head-
rick and Goeden 1990a, 1995), initiation
and termination of mating was simple and
direct, involving little or no pre- or post-
copulatory behaviors. Copulatory postures
exhibited by both sexes were similar to
those generally described for other nonfru-
givorous tephritids (Fig. 5H, I). Five pairs
mated 11 times on successive days for an
average duration of 89 + 14 (range, 22-—
145) min.
Seasonal history.—Like Neaspilota viri-
descens Quisenberry (Goeden and Headrick
1992), Urophora timberlakei Blanc and
Foote (Goeden and Teerink 1995a), and
other Neaspilota and Urophora spp. studied
in southern California (our unpublished
data), the life cycle of X. dichromata fol-
lows the aggregative pattern in which over-
wintering is largely by prepuparia in dead
flower heads. These overwintered prepu-
paria pupariate and emerge as adults the
605
next spring or summer, or Overwintering is
by long-lived adults in reproductive dia-
pause, that return to and aggregate on pre-
blossom host plants during the following
spring and summer to mate and reproduce
(Headrick and Goeden 1995). It is also pos-
sible that adults of X. dichromata that
emerge from flower heads in late summer
(August) produce a second generation in
late-forming, preblossom flower heads of
Hieracium albiflorum, or different, as yet
undetermined, alternate host plant(s), at
higher elevations (Goeden and Headrick
1992; Goeden et al. 1995a).
Natural enemies.—Two species of soli-
tary, hymenopterous, parasitoids were
reared from puparia of X. dichromata in
flower heads of H. albiflorum and were
identified for us as Pteromalus sp. (Pter-
omalidae) and Braconinae sp. (Braconidae).
The former species was the more common
parasitoid.
Biological control.—In the northwestern
United States, three species of Hieracium
native to Eurasia are weeds of forests, pas-
tures, meadows, and wetlands: mouseear
hawkweed (H. pilosella L.), orange hawk-
weed (H. aurantiacum L.), and yellow
hawkweed (H. pratense Tausch). In addi-
tion, two weedy native species, narrowleaf
hawkweed (H. umbellatum L.) and Cana-
dian hawkweed (H. canadense Michaux)
are found in the Northwest (Birdsall and
Quimby 1996). In New Zealand, four intro-
duced species of European origins (includ-
ing H. pilosella) are invasive weeds affect-
ing livestock production and native-plant
and soil conservation in grasslands (Syrett
and Sarospataki 1993). Holm et al. (1979)
listed Canada, Russia, and Finland as ad-
ditional countries in which hawkweeds are
problems. A consortium reportedly has
been formed to identify and assess candi-
date agents for biological control of hawk-
weeds. This group includes representatives
from the U.S. Department of Agriculture,
Agricultural Research Service; University
of Idaho; Agriculture Canada; the Interna-
tional Institute of Biological Control, Swit-
606
zerland and United Kingdom; the Com-
monwealth Scientific and Industrial Re-
search Organization, Australia; and Land-
care Research New Zealand Limited
(Birdsall and Quimby 1996). The final draft
of the present report was given to col-
leagues representing selected member or-
ganizations in this consortium for their pos-
sible interest in X. dichromata as a candi-
date biological control agent, especially for
export to New Zealand, if it will attack and
reproduce on one or more of the weedy spe-
cies introduced there. As noted above, this
nearly monophagous fruit fly attacks only
some native North American species of
Hieracium, and apparently, has not been
reared from any introduced weedy species.
Consequently, it may prove too host spe-
cific for domestic redistribution and aug-
mentation, or otherwise unsuitable for ex-
port, like other native North American in-
sects found attacking native and introduced
weeds (cf., Goeden and Kok 1986, Goeden
and Teerink 1993, Goeden and Palmer
1995)
ACKNOWLEDGMENTS
Once again we sincerely thank A. C.
Sanders, Curator of the Herbarium, Depart-
ment of Botany and Plant Sciences, Uni-
versity of California, Riverside, for identi-
fication of plants mentioned in this paper.
The parasitoids were identified by Michael
Gates, Department of Entomology, Univer-
sity of California, Riverside, and Harry E.
Andersen, Huntington Beach, CA. We also
are grateful to E L. Blanc, D. H. Headrick,
J. B. Johnson, J. McCaffrey, A. L. Norr-
bom, Pauline Syrett, and Linda Wilson for
their helpful comments on or additions to
earlier drafts of this paper.
LITERATURE CITED
Benjamin, F H. 1934. Descriptions of some native
trypetid flies with notes on their habits. U. S. De-
partment of Agriculture Technical Bulletin 401,
95 pp.
Birdsall, J and P. C. Quimby, Jr. 1996. Hawkweeds,
8 pp. In Rees, N., P. C. Quimby, Jr., G. L. Piper,
E. M. Coombs, C. E. Turner, N. R. Spencer, and
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
L. V. Knudson, eds. Biological Control of Weeds
in the West. Western Weed Science Society, Boze-
man, Montana.
Doane, R. W. 1899. Notes on Trypetidae with descrip-
tions of new species. Journal of the New York
Entomological Society 7: 177—193, pl.
Dodson, G. and S. B. George. 1986. Examination of
two morphs of gall-forming Aciurina (Diptera: Te-
phritidae). Ecological and genetic evidence for
species. Biological Journal of the Linnean Society
29: 63-79.
Foote, R. H. and E L. Blanc. 1963. The fruit flies or
Tephritidae of California. Bulletin of the Califor-
nia Insect Survey 7, 117 pp.
1979. New species of Tephritidae from the
United States, Mexico, and Guatemala, with re-
visionary notes. Pan-Pacific Entomologist 55:
161-179.
Foote, R. H., EF L. Blanc, and A. L. Norrbom. 1993.
Handbook of the Fruit Flies (Diptera: Tephritidae)
of America North of Mexico. Cornell University
Press, Ithaca, New York.
Goeden, R. D. 1988. Life history of Trupanea imper-
fecta (Coquillett) on Bebbia juncea (Bentham)
Greene in the Colorado Desert of southern Cali-
fornia (Diptera: Tephritidae). Pan-Pacific Ento-
mologist 64: 345-351.
1990a. Life history of Eutreta diana (Osten
Sacken) on Artemisia tridentata Nuttall in south-
ern California (Diptera: Tephritidae). Pan-Pacific
Entomologist 66: 24-32.
1990b. Notes on the life history of Eutreta
simplex Thomas on Artemisia ludoviciana Nuttall
in southern California (Diptera: Tephritidae). Pan-
Pacific Entomologist 66: 33-38.
Goeden, R. D. and D. H. Headrick. 1991. Notes on
the biology, hosts, and immature stages of To-
moplagia cressoni Aczél in southern California.
Proceedings of the Entomological Society of
Washington 93: 549-558.
1992. Life history and descriptions of im-
mature stages of Neaspilota viridescens Quisen-
berry (Diptera: Tephritidae) on native Asteraceae
in southern California. Proceedings of the Ento-
mological Society of Washington 94: 59-77.
Goeden, R. D., D. H. Headrick, and J. A. Teerink.
1993. Life history and descriptions of immature
stages of Tephritis arizonaensis Quisenberry
(Diptera: Tephritidae) on Baccharis sarothroides
Gray in southern California. Proceedings of the
Entomological Society of Washington 95: 210—
222%
. 1994a. Life History and descriptions of im-
mature stages of Procecidochares flavipes Aldrich
(Diptera: Tephritidae) on Brickellia spp. in south-
ern California. Proceedings of the Entomological
Society of Washington 96: 288-300.
1994b. Life history and description of im-
VOLUME 99, NUMBER 4
mature stages of Paroxyna genalis (Thomson)
(Diptera: Tephritidae) on native Asteraceae in
southern California. Proceedings of the Entomo-
logical Society of Washington 96: 612-629.
1995a. Life history and description of im-
mature stages of Urophora timberlakei Blanc and
Foote (Diptera: Tephritidae) on native Asteraceae
in southern California. Proceedings of the Ento-
mological Society of Washington 97: 780-791.
1995b. Life history and descriptions of im-
mature stages of Valentibulla californica (Coquil-
lett) (Diptera: Tephritidae) on Chrysothamnus
nauseosus (Pallas) Britton in southern California.
Proceedings of the Entomological Society of
Washington 97: 548-560.
Goeden, R. D. and L. T. Kok. 1986. Comments on a
proposed “new” approach for selecting agents for
the biological control of weeds. Canadian Ento-
mologist 118: 39-47.
Goeden, R. D. and W. A. Palmer. 1995. Lessons
learned from studies of the insects associated with
Ambrosiinae in North America in relation to the
biological control of weedy members of this
group, pp. 565-573. Jn Delfosse, E. S., and R. R.
Scott, eds. Proceedings Eighth International Sym-
posium on Biological Control of Weeds, Lincoln
University, Canterbury, New Zealand, 2—7 Feb-
ruary, 1992. Department of Scientific and Indus-
trial Research, New Zealand; Commonwealth Sci-
entific and Industrial Research Organization, Mel-
bourne, Australia.
Goeden, R. D. and J. A. Teerink. 1993. Phytophagous
insect faunas of Dicoria canescens and Iva axil-
laris, native relatives of ragweeds, Ambrosia spp.,
in southern California, with analyses of insect as-
sociates of Ambrosiinae. Annals of the Entomo-
logical Society of America 86: 37—50.
1996a. Life histories and descriptions of
adults and immature stages of two cryptic species,
Aciurina ferrugenia (Doane) and A. michaeli new
species (Diptera: Tephritidae) on Chrysothamnus
viscidiflorus (Hooker) Nuttall in southern Califor-
nia. Proceedings of the Entomological Society of
Washington 98: 415-438.
. 1996b. Life history and descriptions of adults
and immature stages of Aciurina idahoensis
Steyskal (Diptera: Tephritidae) on Chrysothamnus
viscidiflorus (Hooker) Nuttall in southern Califor-
nia. Proceedings of the Entomological Society of
Washington 98: 681-694.
. 1996c. Life history and descriptions of adults
and immature stages of Aciurina semilucida
(Bates) (Diptera: Tephritidae) on Chrysothamnus
viscidiflorus (Hooker) Nuttall in southern Califor-
nia. Proceedings of the Entomological Society of
Washington 98: 752-766.
. 1997. Notes on life histories and descriptions
607
of adults and immature stages of Procecidochares
kristineae and P. lisae new spp. (Diptera: Tephrit-
idae) on Ambrosia spp. in southern California.
Proceedings of the Entomological Society of
Washington 99: 67-88.
Headrick, D. H. and R. D. Goeden. 1990a. Descrip-
tion of the immature stages of Paracantha gentilis
(Diptera: Tephritidae). Annals of the Entomolog-
ical Society of America 83: 220-229.
. 1990b. Resource utilization by larvae of Par-
acantha gentilis (Diptera: Tephritidae) in capitula
of Cirsium californicum and C. proteanum (As-
teraceae) in southern California. Proceedings of
the Entomological Society of Washington 92:
512-520.
1991. Life history of Trupanea californica
Malloch (Diptera: Tephritidae) on Gnaphalium
spp. in southern California. Proceedings of the En-
tomological Society of Washington 93: 559-570.
1994. Reproductive behavior of California
fruit flies and the classification and evolution of
Tephritidae (Diptera) mating systems, Studia Dip-
terologica 1(2): 194—252.
Headrick, D. H., R. D. Goeden, and J. A. Teerink.
1996. Life history and description of immature
stages of Dioxyna picciola (Bigot) (Diptera: Te-
phritidae) on Coreopsis spp. (Asteraceae) in
southern California. Proceedings of the Entomo-
logical Society of Washington 98: 332-349.
Holm, Je Ve Rancho, Jo Py Herberger, andeD>
Plucknett. 1979. A Geographical Atlas of World
Weeds. John Wiley & Sons, New York.
Munz, P. A. 1974. A Flora of Southern California.
University of California Press, Berkeley.
Norrbom, A. L., L. E. Carroll, and A. Freidberg. 1997.
Status of Knowledge. Jn Thompson, F. C., ed.
Fruit Fly Expert System and Biosystematic Infor-
mation Database, Myia and Diptera Data Dissem-
ination Disk (CD-ROM).
Novak, J. A., W. B. Stolzfus, E. J. Allen, and B. A.
Foote. 1967. New host records for North Amer-
ican fruit flies (Diptera: Tephritidae). Proceedings
of the Entomological Society of Washington 69:
146-148.
Snow, W. A. 1894. Descriptions of North American
Trypetidae, with notes. Kansas Entomological So-
ciety Quarterly 2: 159-174.
Steck, G. J. and R. A. Wharton. 1986. Descriptions
of immature stages of Eutreta (Diptera: Tephriti-
dae). Journal of the Kansas Entomological Society
59(2): 296-302.
Syrett, P. and M. Sarospataki. 1993. Prospects for bi-
ological control of hawkweeds with insects. pp.
426-432. In Prestidge, R. A. (ed.). Proceedings
6th Australasian Grassland Inverterbrate Ecology
Conference, AgResearch, Hamilton, New Zea-
land.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 608-613
A NEW SPECIES OF MEGASTIGMUS DALMAN (HYMENOPTERA:
TORYMIDAE) REARED FROM SEEDS OF ATLANTIC WHITE CEDAR
(CUPRESSACEAE), WITH NOTES ON INFESTATION RATES
J. J. TURGEON, K. KAMIJO, AND G. DEBARR
(JJT) Canadian Forest Service, Great Lakes Forestry Centre, PO. Box 490, Sault Ste.
Marie, Ontario, Canada P6A 5M7; (KK) Nishi 2 Minami 6, Bibai, Hokkaido, 072 Japan;
(GD) U. S. Department of Agriculture, Forest Service, Southern Research Station, 320
Green St., Athens, GA 30602-2044, U.S.A.
Abstract.—A new species, Megastigmus thyoides Kamijo (Hymenoptera: Torymidae),
which emerged from seeds of Atlantic white cedar, Chamaecyparis thyoides (L.) B.S.P.,
collected in eastern United States is described and illustrated. This is the first record of
this genus exploiting seeds of Cupressaceae in the Nearctic region. An average of 7% of
the seeds collected from five sites in North Carolina were infested with this seed chalcid.
Key Words:
For several tree species, insects often
represent the most important source of seed
mortality during the pre-dispersal phase of
seed development (1.e., while still devel-
oping on the tree). Most insects exploiting
the seed cones of commercially important
conifers native to North America have al-
ready been identified, as surveys over the
past 30 years have focused almost entirely
on these trees (Miller et al. 1995, Turgeon
et al. 1994). Conversely, knowledge of the
entomofauna of native conifers that are of
lesser economic importance is scarce. Dur-
ing a survey initiated in 1994 to address
some of these knowledge gaps, we discov-
ered specimens of an undescribed species
of Megastigmus Dalman (Hymenoptera:
Torymidae) feeding in seeds of Atlantic
white cedar, Chamaecyparis thyoides (L.)
B.S.P. (Cupressaceae). This species is de-
scribed below and the rates of seed infes-
tation observed are presented.
There are approximately 100 described
species of Megastigmus worldwide (Bou-
Hymenoptera, Torymidae, Megastigmus thyoides, new species, Chamae-
cyparis thyoides, seed chalcid
éek 1988, Xu and He 1995). Most species
are phytophagous (Roux and Roques 1996);
exploiting seeds of either coniferous or de-
ciduous trees or shrubs (Milliron 1949;
Boucek 1988). Conifers from the Pinaceae,
Cupressaceae and Taxodiaceae families are
exploited by at least 45 species of Megas-
tigmus (Yates 1986, Turgeon et al. 1994, Xu
and He 1995, A. Roques, personal com-
munication).
Morphological terminology and mea-
surements generally follow those of Boucek
(1988). The following abbreviations are
used throughout the text: FI—F7 = funicu-
lar segments 1-7; T1-T7 = tergites 1-7.
Megastigmus thyoides Kamijo,
NEW SPECIES
(Figs. 1—4)
Female.—Body length 1.4 to 1.7 mm.
Body brownish yellow: T1—T4 each with
vague dark band. Antenna with pedicellus
and flagellum dark brown. Wings subhya-
VOLUME 99, NUMBER 4
609
Figs. 1-4. Megastigmus thyoides. 1, Frontal view of female head and antenna. 2, Lateral view of female. 3,
Forewing of female. 4, Stigmal vein of forewing of male.
line; veins including basal vein dark brown.
Ovipositor sheath blackish.
Head in dorsal view about 1.6 times as
broad as long, with occiput weakly emar-
ginate and temples roundly converging.
POL nearly 1.6 times OOL. Vertex weakly
strigate; ocellar area, frons and lower face
strigate-reticulate. Head covered with
sparse, black setae, but most setae on lower
face whitish. Eyes separated by 1.6 times
their height. Malar space slightly longer
than half height of eye. Antennal scrobes
almost reaching median ocellus; toruli sit-
uated exactly in middle of height of head
(Fig. 1). Scape reaching level of vertex;
combined length of pedicellus and flagel-
lum 1.2 times breath of head; flagellum
weakly clavate; anellus slightly transverse
to quadrate; F1 nearly as long as pedicellus;
F7 quadrate to slightly transverse; sensilla
disposed in 1 row on each funicle segment.
Thorax strongly arched in lateral view
(Fig. 2), 1.5 to 1.6 times as long as broad
in dorsal view. Pronotum nearly 1.6 times
as broad as long, strigate-reticulate. Meso-
scutum a little longer than pronotum; mid-
610
lobe with distinct, raised reticulation, and 2
or 3 black setae present on each side; scap-
ulae reticulate. Scutellum as long as broad,
sculptured like mesoscutal mid-lobe, with 3
or 4 setae on each side; frenum more
coarsely reticulate, with frenal line indis-
tinct, almost straight. Propodeum with
strongly raised reticulation, sometimes me-
dian carina present; callus with 3 or 4 whit-
ish setae. Prepectus with oblique carina
rather indistinct. Mesepisternum and mes-
epimeron densely reticulate.
Forewing: marginal vein slightly shorter
than postmarginal; stigmal vein a little lon-
ger than half length of marginal, with stig-
ma slender, basal cell with 1 or 2 apical
setae, open in basal half; basal vein distinct-
ly pigmented; speculum rather large, closed
below (Fig. 3).
Gaster a little longer than thorax; hind
margins of Tl and T2 barely incised me-
dially. Ovipositor sheath weakly clavate,
about as long as thorax or 1.6 to 1.9 times
as long as hind tibia (Fig. 2).
Male.—Differs from female as follows.
Body length 1.4 to 1.9 mm. Ocellar area usu-
ally dark brown; scutellum posteriorly, meta-
notum and propodeum sometimes darker;
gaster with T1I—-T5 more extensively dark
brown. Head in dorsal view 1.6 to 1.7 times
as broad as long; combined length of pedi-
cellus and flagellum nearly 1.4 times breadth
of head; flagellum filiform, covered with
denser setae; F7 quadrate to slightly longer
than broad. Thorax more slender, sometimes
nearly 1.8 times as long as broad. Mid-lobe
of mesoscutum with 3 to 5 setae on each side.
Scutellum slightly longer than broad. Fore-
wing with stigma slightly broader; basal cell
with 2 or 4 setae, open basally (Fig. 4). Gas-
ter much longer than thorax.
Types.—Holotype 2, Dare County, North
Carolina, USA, ex seed of Chamaecyparis
thyoides collected in 1994, deposited in Ca-
nadian National Collection of Insects (CNCI).
Paratypes: 14 2, 15 6d, with same rearing
data as holotype. 2 d and 2 2 paratypes de-
posited in the National Museum of Natural
History, Smithsonian Institution, and in the
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
personal collection of JJT at the Canadian
Forest Service-Great Lakes Forestry Centre,
Sault Ste. Marie, Ontario, Canada. Another
pair is at Hokkaido University, Sapporo, Ja-
pan. The remainder of the paratypes are de-
posited in the CNCI.
Distribution.—Dare, Wayne and Perquim-
ins Counties, North Carolina, U.S.A.
Host.—Seeds of Atlantic white cedar,
Chamaecyparis thyoides (L.) B.S.P. (Cu-
pressaceae).
Discussion.—Megastigmus thyoides
Kamijo is the first record of this genus ex-
ploiting seeds of Cupressaceae in the Ne-
arctic Region. In the Palaearctic Region
however, seeds of Cupressaceae are host to
thirteen species of Megastigmus. Eight of
these thirteen species infest seeds of Junip-
erus spp. Seeds of Cupressus spp., Cha-
maecyparis spp. and Thujopsis spp. are ex-
ploited by three, one and one species of
seed chalcid, respectively (Zerova and Ser-
yogina 1994, Xu and He 1995). There is
also a record of this genus exploiting seeds
of Cupressaceae in the Oriental Region,
where M. cupressi Mathur infests seeds of
Cupressus torulosa Don. (Mathur 1955).
Megastigmus thyoides Kamijo is charac-
terized by the reticulate sculpture on the
head and thorax, the elongate stigma, the
short ovipositor, and the strongly arched
thorax in longitudinal axis. This species ap-
pears closely related to two phytophagous
species from Japan, M. chamaecyparidis
Kamijo and M. thuyopsis Yano that exploit
seeds of Chamaecyparis obtusa (Sieb. and
Zucc.) Endl. and Thujopsis dolabrata Sieb.
and Zucc. (Cupressaceae), respectively
(Kamijo 1962). All three species have a
row of sensilla on all funicular segments, a
brownish-yellow body, and the pronotum,
mesoscutum and scutellum weakly sculp-
tured with a small number of black bristles.
However, unlike M. thyoides, the two Jap-
anese species have the head, pronotum and
mesoscutum weakly strigate, a weakly
arched thorax, nearly 1.9 times as long as
broad in dorsal view, a stigma less slender,
and ovipositor sheaths 1.7 times as long as
VOLUME 99, NUMBER 4
thorax in M. chamaecyparidis, and 1.3
times as long in M. thuyopsis.
Megastigmus thyoides also resembles the
Nearctic species M. hoffmeyeri Walley,
which feeds in seeds of Tsuga canadensis
(L.) Carr. (Pinaceae) (Milliron 1949) and
the Japanese M. tsugaphilus Kamijo (Ka-
mijo 1962). Both M. thyoides and M. hoff-
meyeri have a body entirely brownish yel-
low and an elongate stigma, but according
to the redescription given by Milliron
(1949), M. hoffmeyeri has a longer ovipos-
itor, pronotum and mesoscutum rugulose,
and distal funicle segments longer than M.
thyoides. Megastigmus tsugaphilus also has
a brownish body and a strongly arched tho-
rax which resembles that of M. thyoides. In
M. tsugaphilus, however, besides the long
Ovipositor and the strigate sculpture on the
head and thorax, the funicular segments
have two rows of sensilla, the basal cell of
the forewing has many setae, and the mid-
lobe of scutellum usually has five setae on
each side (Kamijo 1962).
The most recent key for the Nearctic spe-
cies of Megastigmus is that by Milliron
(1949). Females of M. thyoides key best at
a slightly modified (in bold) couplet 3 of
Milliron’s key.
3. Pronotum weakly arched above and without
sharp transverse rugae; mid-lobe of meso-
scutum with fine, reticulate sculpture; stigma
narrow-elongate, pointed or rounded on
lower margin; small species (1.2 to 2.2 mm.)
Ee eee let ie et a een a aie 3a
— Pronotum more strongly arched and usually
with at least one distinct transverse ruga; mid-
lobe of mesoscutum transversely rugulose
or sometimes shingled; stigma more or less
Ovalfor circular sit. ate on eee eee eee 5)
3a. Stigma narrow-elongate oval with lower mar-
gin rounded (Fig. 3); setae on head and tho-
AGS GorrwioN WIE oon co aoc thyoides Kamijo
— Stigma surrounded by cloud or somewhat tri-
angular, with lower margin pointed (Milliron
1949: Figs. 7, 9); setae on head and thoracic
dorsum pale
Males of M. thyoides key out at couplet
43. Some characters of M. thyoides used in
611
the following modified couplet are not in-
cluded in our description (e.g., length of
hind legs).
43. Hind tarsus distinctly greater than two-thirds
of the hind tibia; proximal tarsal segments
rather long, the dorsal measurement of the
first about | *% that of the second ......
— Hind tarsus nearer to two-thirds as long as
the hind tibia; proximal segments of hind
tarsus comparatively shorter than above .. 44
Pronotum nearly as long as wide; mesoscu-
43a
43a.
tum and scutellum transversely rugulose;
head and thorax brownish yellow with
blackishtmarkinese sess ee: rafni Hoffmeyer
— Pronotum distinctly wider than long; meso-
scutum and scutellum densely reticulate;
head and thorax entirely brownish yellow
2) vg spits) chock cad ana oaceecd <a aac’ thyoides Kamijo
Etymology.—This species is named after
its host, C. thyoides.
Biology.—This species is a phytopha-
gous insect developing in seeds of its host.
Most species of this genus developing in
conifer seeds have a univoltine life cycle,
Overwintering as a 5th instar in seed on the
ground (Hedlin et al. 1980). Thus far, only
M. spermotrophus Wachtl has been shown
to oviposit and develop in unfertilized seeds
(Niwa and Overhulser 1992, Rappaport et
al. 1993). Whether this species also devel-
ops in unfertilized seeds of C. thyoides will
be investigated together with its potential
for prolonged diapause, a mechanism used
by several species of this genus to compen-
sate for annual fluctuations in cone abun-
dance (Roux 1995).
Seed cones of C. thyoides were collected
in late-October early-November 1994 from
various sites (each identified with a different
seed lot number) in North Carolina (Table
1). Cones were dried at room temperature
and the seeds extracted. A subsample of
seeds from each lot was spread as uniformly
as possible on a sheet of Kodak Industrex
620 X-ray paper and irradiated with a Hew-
lett Packard Faxitron 43805N X-Ray system
for 30 sec at 15 Kv and 3 mA. The exposed
paper was developed using a model P-1 Ko-
612 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Quality of C. thyoides seeds collected in North Carolina together with infestation rates by M.
thyoides.
Infested
by
Filled Empty M. thy-
Collection Site County Seed Lot Number Type of Stand (%) (%) oides
Sasser Tract (Goldsboro) Wayne WC-NC 94-6A 5-y-old tree farm 37.7 58.5 3.8
Sasser Tract (Goldsboro) Wayne WC-NC 94-6B 5-y-old tree farm 28.0 67.4 4.6
Milltown Road (East Lane) Dare WCNC 94-7 Natural stand 126) VSIe2 9 Vilee
D-7 (Elizabeth City) Perquimins WCNC 94-8 Plantation 35 eGSe2. let
Dare Bomb Range (Stumpy Point) Dare WCNC 94-9 Stand-logged L0:0° 75:0) sa
Average 23.8 6910 V2
dak Industrex Instant Processor. On each ra-
diograph an acetate overlay with 1 cm grid
intervals and coordinates from a random
number table were used to locate the centre
points for 10, non-overlapping circles (diam.
of 3 cm). A 10x hand lens was used to ex-
amine the radiographic images and deter-
mine for each seed lot the numbers of filled,
empty and chalcid-infested seeds within
each circle.
The proportions of filled, empty and
chalcid-infested seeds varied substantially
among seed lots (Table 1). On average, al-
most 70% of the seeds from each seed lot
were empty. The true impact, as defined by
Rappaport et al. (1993), of M. thyoides
could not be assessed because the number
of sound seeds in chalcid-excluded cones
could not be determined. Nonetheless, seed
infestation rates by M. thyoides appear rel-
atively low, never exceeding 15% in a seed
lot. These rates are similar to those reported
for other species of Megastigmus exploiting
conifers in their native range (Hedlin et al.
1980, Roques 1983).
ACKNOWLEDGMENTS
We thank Dr. J. Huber of the CNC for
assistance in critical point drying and
mounting the valuable specimens, for com-
ments on an earlier draft of this manuscript,
and for assisting with the key; the North
Carolina Forest Service for collecting seed
cones; and, K. Barber and K. Nystrom for
their valuable suggestions on an earlier
draft.
LITERATURE CITED
Boucek, Z. 1988. Australasian Chalcidoidae (Hyme-
noptera). A biosystematic revision of genera of
fourteen families, with a reclassification of spe-
cies. CAB International Institute of Entomology,
Wallingford, England. 832 pp.
Hedlin, A. FE, H. O. Yates II, D. Cibrian-Tovar, B. H.
Ebel, T; W. Koerber, and E. P. Merkel. 1980.
Cone and seed insects of North American coni-
fers. Environment Canada, Canadian Forest Ser-
vice, Ottawa/ USDA Forest Service Washington/
Secretaria de Agricultura y Recursos Hidraulicos,
Mexico. 122 pp.
Kamijo, K. 1962. A revision of the species of the
Megastigminae occurring in Japan (Hymenoptera:
Chalcidoidea). Insecta Matsumurana 25: 18—40.
Mathur, R. N. 1955. A new species of Megastigmus
Dalman (Hymenoptera: Chalcidoidea) from the
seeds of Cupressus torulosa Don. Proceedings of
the Royal Entomological Society of London. (B)
24: 91-94.
Miller, G. E., J. J. Turgeon, and P. de Groot. 1995.
Cone and seed insects, pp. 293-311. In Arm-
strong, J. A. and W. G. H. Ives, eds. Forest Insect
Pests in Canada, Natural Resources Canada: Ot-
tawa.
Milliron, H. E. 1949. Taxonomic and biological in-
vestigations in the genus Megastigmus, with par-
ticular reference to the taxonomy of the Nearctic
species (Hymenoptera: Chalcidoidea: Callimomi-
dae). The American Midland Naturalist 41: 257—
420.
Niwa, C. G. and D. L. Overhulser. 1992. Oviposition
and development of Megastigmus spermotrophus
(Hymenoptera: Torymidae) in unfertilized Doug-
las-fir seed. Journal of Economic Entomology 85:
2323-2328.
Rappaport, N., S. Mori, and A. Roques. 1993. Esti-
mating effect of Megastigmus spermotrophus
(Hymenoptera: Torymidae) on Douglas-fir seed
production: the new paradigm. Journal of Eco-
nomic Entomology 86: 845-849.
VOLUME 99, NUMBER 4
Roques, A. 1983. Les insects ravageurs des cOnes et
graines de coniféres en France. INRA: Paris. 135
PP- )
Roux, G. 1995. Etude comparative des mécanismes
d’induction de la diapause prolongée chez quel-
ques insectes spécifiques des cénes de coniféres.
Ph.D. Thesis, Université d’Orléans, France.
Roux, G. and A. Roques. 1996. Biochemical genetic
differentiation among seed chalcid species of ge-
nus Megastigmus (Hymenoptera: Torymidae). Ex-
perientia 52: 522-530.
Turgeon, J. J.. A. Roques, and P. de Groot. 1994. In-
sect fauna of coniferous seed cones: diversity, host
613
plant interactions and management. Annual Re-
view of Entomology 39: 179-212.
Xu Z. H. and He J. H. 1995. Note on species of phy-
tophagous group of Megastigmus (Hymenoptera:
Torymidae) from China. Entomotaxonomia 17:
243-253. [In chinese with english abstract.]
Yates, H. O. Il. 1986. Checklist of insect and mite
species attacking cones and seeds of world coni-
fers. Journal of Entomological Science 21: 142—
168.
Zerova, M. D. and L. Ya. Seryogina. 1994. The seed-
eating Chalcidoidea of Palaearctics. National
Academy of Sciences of Ukraine. 237 pp. [In Rus-
sian. }
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 614-622
SHOOT OR STAB? MORPHOLOGICAL EVIDENCE ON THE UNRESOLVED
OVIPOSITION TECHNIQUE IN STYLOGASTER MACQUART (DIPTERA:
CONOPIDAE), INCLUDING DISCUSSION OF BEHAVIORAL
OBSERVATIONS
MARION KOTRBA
Department of Entomology, Smithsonian Institution, National Museum of Natural His-
tory NHB 169, Washington, DC 20560, U.S.A.
Abstract.—Behavioral observations on the oviposition behavior of Stylogaster are re-
viewed in combination with data on the egg morphology, egg placement, and the female
postabdomen. The internal female reproductive tract of S. stylosa Townsend is described.
While some behavioral observations seem to imply a shooting oviposition technique, the
morphological evidence favors stabbing oviposition.
Key Words:
eggs
In 1961 Rettenmeyer described the ovi-
position behavior of Stylogaster (Diptera:
Conopidae) above swarms of raiding army
ants (Hymenoptera: Formicidae, Ecitoni-
nae) in Panama. “‘While hovering 30 to 70
cm above the ground, females of Stylogas-
ter characteristically flipped their abdomens
down, or their whole body shifted down-
ward a few centimeters. This movement
suggested that the flies were dropping or
shooting eggs at the ants below” (p. 1004,
emphasis mine). He further reported that on
several occasions Stylogaster was seen to
“dart after cockroaches.”’
Several other authors have also observed
hovering Stylogaster above swarms of army
ants in Central America (Fig. 1) or Africa:
‘“‘occasionally one of the flies darting with
great quickness toward the ground”’ (Bates
1863: 366); “‘now and again darting at
them, without doubt for the purpose of ovi-
positing in their bodies”’ (Townsend 1897:
23); “suddenly pouncing down (apparently
for the purpose of ovipositing)”’ upon flee-
ing cockroaches (Carpenter 1915, cited in
Female postabdomen, oviscapt, female reproductive tract, spermathecae,
Stuckenberg 1963); or ““pouncing like little
warplanes amidst the ants”? (Lindner 1955:
23, translated from German). Smith & Cun-
ningham-van Someren (1985) observed that
small cockroaches escaping from raiding
army ants were attacked by Stylogaster.
The closest observation was published by
Lopes (1937: 260), who observed that some
Stylogaster females approached an orthop-
teran and ‘“‘with an extremely rapid move-
ment inserted an egg in the abdomen be-
tween the tergites”’ (translated from Portu-
guese). Of the many documented observa-
tions none describes contact, grasping, or
seizing of the victim.
The actual process of oviposition, 1.e.,
how the egg is inserted into the host’s body,
remains obscure. Based on the investigation
of collected material, Smith (1967: 52)
speculated that “‘the eggs are probably laid
by a stabbing movement of the abdomen,
with the oviscapt thrust forward between
the legs.’’ He further surmised that “‘the
hovering females frequently flick the tips of
their abdomens rapidly, but it is doubtful if
VOLUME 99, NUMBER 4
Fig. 1. Hovering Stylogaster female, Los Alturas,
Costa Rica.
Oviposition accompanies this movement,
since considerable force at close quarters
must surely be needed for the egg to pen-
etrate the host cuticle.”’
The present study follows up on this
problem. Could it be, that (A) Stylogaster
literally shoots eggs at its host, as suggested
by the flipping movements of the abdomen
(Rettenmeyer 1961) and the fact that no
physical contact with the host has been re-
ported? Or (B) are the eggs stabbed into the
host’s integument, as suggested by Smith
(1967, 1979)? Or (C) are the eggs inserted
more conventionally after the victim has
been seized, which has so far escaped ob-
servation? The available information on the
female postabdomen of Stylogaster, its
dart-like eggs, and their random placement
on the host’s body is reviewed and supple-
mented by a description of the internal fe-
male reproductive tract of S. stylosa Town-
send. The possibilities for shooting or stab-
615
bing oviposition in Stylogaster are weighed
and approaches for further testing are sug-
gested.
MATERIALS AND METHODS
The description of the internal female re-
productive system of Stylogaster is chiefly
based upon two specimens of S. stylosa,
which were collected in Costa Rica and dis-
sected immediately after killing. The female
reproductive tract was removed, placed on
a glass slide, and embedded in Polyvinyl-
lactophenol with an admixture of Chlorazol
E (Streng 1976). This medium slowly mac-
erates the tissue while the dye is enriched
in unsclerotized cuticle, staining it blue.
The terminalia of S. frauci Smith were dis-
sected while held in lactic acid. The prep-
arations were investigated and documented
with an Olympus BX50 compound micro-
scope equipped with a drawing tube and
Olympus SC35 camera, and with a Wild M
400 Photomakroskop.
RESULTS
Female Postabdomen
The female postabdomen of Stylogaster
has been depicted for various species
(Lopes 1937, 1938, 1971, Lopes and Mon-
teiro 1959, Smith 1967, Smith and Peterson
1987). It is characterized by a tubular ovi-
scapt (Smith 1966), which can be extremely
long and narrow in some species (Figs. 2,
3). In the Brazilian S. macrura Lopes the
oviscapt is twice as long as the remainder
of the body excluding the antennae (Lopes
1938). The terminalia of Stylogaster lack
any structures able to penetrate the host in-
tegument, as found in other Conopidae
(Howell 1967, Smith & Peterson 1987) and
other parasitic flies such as Pipunculidae
(Hardy 1987) and Pyrgotidae (Steyskal
1987). Instead, the vulva is surrounded by
various elongate structures (“egg guides”’
Smith 1979). The cerci are long and slen-
der, the subanal plate is elongate. Sternum
8 ends in a pair of pointed lobes and bears
a pair of long and slender lateral append-
616
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 2-5. Stylogaster frauci female. 2, Total specimen in alcohol. 3, Abdomen cleared in lactic acid; eggs
and spermathecae visible within the abdomen; one egg protruding from the vulva. 4, Terminalia. 5, Terminalia
of with egg lodged in terminal chamber.
ages with inward curved endlobes (Figs. 4—
2h
As reported above, Stylogaster females
frequently “‘flip’’ the abdomen while in
flight, rapidly flexing it under the body so
that the oviscapt is directed anteroventrally.
This movement is facilitated by the absence
of sclerotized sternites in the first five ab-
dominal segments (Stuckenberg 1963). The
resulting flexed posture is often seen in
pinned or preserved specimens (Fig. 2).
Egg
The eggs of Stylogaster show ‘‘remark-
able adaptations for the penetration of, and
retention in, the host cuticle’? (Smith 1966:
264). They have been depicted for various
species (Lopes 1937, 1938, 1971 Lopes and
Monteiro 1959, Stuckenberg 1963, Smith
1967, 1979, Hinton 1981, Smith and Cun-
ningham-van Someren 1985). In general
Stylogaster eggs are elongate with a trans-
lucent brown chorion. The posterior end is
black due to heavy sclerotization, sharply
pointed, and bears up to four recurrent
spines (Figs. 5, 12). Moreover, a membra-
nous bladder can be extruded from an area
near the posterior end. This bladder is be-
lieved to be everted by osmosis after the
egg is inserted into the host, serving to an-
chor the egg and erect the recurrent spines
(Rettenmeyer 1961, Stuckenberg 1963).
Stylogaster eggs are very different from
those of other conopids, which have a thin
whitish chorion, bear appendages at the an-
terior egg pole in the vicinity of the micro-
pyle, and lack an extrusible bladder (de
Meijere 1904, Smith and Peterson 1987).
Egg Placement on the Host Body
The accuracy of Stylogaster oviposition
resembles that of a faulty shotgun. Stuck-
enberg (1963) and Smith (1967, 1969)
found Stylogaster eggs to be positioned ran-
domly on head, thorax, and abdomen of
various muscids in South Africa, a few
even in the wings. Stuckenberg (1963: 258)
explained Rettenmeyer’s (1961) findings of
a nonrandom distribution of Stylogaster
eggs on tachinids of the genus Calodexia
VOLUME 99, NUMBER 4
Figs. 6—9. Terminalia of Stylogaster frauci. 6, Ven-
tral view. 7, View from upper left. 8, Lateral view
(compare Fig. 4). 9, Lateral view with an egg lodged
in the terminal chamber (compare Fig. 5). Abbrevia-
tions: an = anus; ce = cerci; ll = lateral lobes on
sternum eight; sa = subanal plate; s8 = sternum eight;
tc = terminal chamber; vr = scleortized ventral rod.
Wulp from Panama as a result of “‘Stylo-
gaster flies darting after cockroaches which
were also being pursued by a number of
Calodexia. If the tachinids were actively
chasing the orthopterons, they would be
more prone to receive a Stylogaster attack
from behind owing to their forward motion
which would necessitate some following by
the Stylogaster, and also because the ab-
domen would be more susceptible owing to
the attitude of the fly and more exposed as
the wings would be in use.”’
The accuracy of host choice seems like-
617
wise poor. Stylogaster eggs were found not
only on members of the presumed hosts,
entailing certain Orthoptera, Tachinidae,
and silvicolous dung-breeding Muscidae
(Lindner 1955, Rettenmeyer 1961, Stuck-
enberg 1963, Smith 1966, 1969, Smith and
Cunningham-van Someren 1985), but on
various other insects that might have been
hit accidentally (Rettenmeyer 1961, Stuck-
enberg 1963, Smith 1966, Smith and Cun-
ningham-Van Someren 1985). I even found
a Stylogaster egg protruding from the me-
dian occipital sclerite of a Stylogaster sty-
losa female from Costa Rica.
Internal Female Reproductive Tract
The available information on the internal
female reproductive tract of Stylogaster has
been very limited. Lopes (1937) illustrated
the spermathecal ducts of S. stylata (Fabri-
cius) with a detail on the spermathecal duct
pump. Moreover, Smith (1967) sketched the
spermathecae of several African species.
The present study of S. stylosa comple-
ments these data.
The internal female reproductive tract of
Stylogaster stylosa (Fig. 10) comprises
paired ovaries and lateral oviducts, a com-
mon oviduct, a tubular vagina, and a scler-
otized terminal chamber. Into the dorsal
wall of the vagina open two spermathecae,
a third, blind duct between the spermathe-
cae, a pair of accessory glands posterior to
the spermathecae, and two additional,
smaller evaginations, one anterior to the
spermathecae and one posterior to the ac-
cessory glands. A roughly conical evagi-
nation emanates from the anteroventral por-
tion of the vagina.
The ovaries usually contain a large num-
ber of mature and fully sclerotized eggs, but
apparently no earlier egg stages. One of the
S. stylosa specimens contained about 120
mature eggs. This is consistent with Smith’s
(1966, 1967) and Stuckenberg’s (1963)
findings of 60-128 mature eggs in African
Stylogaster. Usually some eggs have al-
ready proceeded into the oviducts (Figs. 3,
2):
618 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
A c
( 10
Fig. 10. Internal female reproductive tract of Stylogaster stylosa. Ovaries, one spermatheca, and one accessory
gland omitted. Abbreviations: ae = anteroventral evagination; ag = accessory glands; ap = accessory gland
duct pump; bd = blind duct; de 1, 2 = dorsal evaginations; Il = lateral lobes; tc = roof of the terminal chamber;
sb = sclerotized base of the spermatheca: sd 1, 2 = apical and basal part of the spermathecal duct; sdp =
spermathecal duct pump; sp = spermatheca; va = vagina; vr = scleortized ventral rod.
VOLUME 99, NUMBER 4
619
Figs. 11—12. Internal female reproductive organs of Stylogaster stylosa. 11, Pump structure in the spermathecal
ducts. 12, Common oviduct containing an egg and apical portion of the vagina. Abbreviations: ae = anteroventral
evagination; de 1 = dorsal evagination; mb = muscle band extending anteriorly from the anteroventral evagi-
nation; sd 1 = apical part of spermathecal ducts; sd 2 =
The two spermathecae are large, round,
and almost entirely membranous. However,
the spermathecal duct insertion at their base
is surrounded by a small sclerotized area
from which black spoke-like ridges radiate
into the membranous part (Fig. 10). The
spermathecal ducts are very long and thin,
and show considerable differentiation along
their length. The basal one-fourth consists
of thick, colorless cuticle surrounded by a
sheath of longitudinal muscles. The apical
three-fourths consists of thinner, translucent
brownish cuticle lacking a muscle sheath.
Between the two portions there is an elab-
orate pump structure, consisting of a thick
cuticular cylinder surrounded by a bulky
layer of spiral muscles (Fig. 11). A third,
blind duct is inserted between the two sper-
mathecal ducts. It is only one-fourth of their
basal part of spermathecal ducts.
length, and its walls consist of thick, multi-
layered rubber-like cuticle. The homology
of this duct is unclear. The accessory glands
have a tubular gland reservoir and mem-
branous, narrow ducts, with a widened
muscular pump region in their apical part.
The anteroventral evagination (Fig. 12) is
thick-walled and surrounded by thick mus-
cles like the rest of the vagina. Moreover,
from its anterior wall a conspicuous muscle
band extends anteriorly along the ventral
side of the common oviduct. The homology
of this anteroventral evagination is uncer-
tain, although its position is reminiscent of
the ventral receptacle of other acalyptrate
Schizophora.
The membranous, tubular part of the va-
gina opens through a valve into a wider
posterior portion, whose ventral wall is re-
620
inforced by a sclerotized rod. This rod con-
nects posteriorly to the dorsal rim of a scler-
otized oval terminal chamber that is invag-
inated from the posterior edge of sternum 8
(Figs. 4, 8, 9). In one of the specimens I
found an egg fitted into this chamber, its
pointed tip protruding from the vulva (Figs.
5, 9). I encountered the same condition in
some dissections of S. currani Aldrich, S.
biannulata Say, and S. frauci. Smith (1967)
similarly figured a S. leonum Westwood
specimen with the black, pointed tip of an
egg protruding from the vulva. In S. frauci
I could establish that the terminal chamber
and surrounding sclerites have the same
shape and position, whether containing an
egg or not (Figs. 4, 5). There is no apparent
evidence of elastic structures that might be
able to store and rapidly release energy for
the launching of an egg.
Dissections of other Stylogaster species
show that the spermathecae, accessory
glands, and additional dorsal evaginations
vary considerably within the genus. These
organs are not regarded in the following
discussion and will be described in detail
elsewhere. The constitution of the ovaries,
the vagina, and the anteroventral evagina-
tion was found to be quite similar across
the species, including African and Austra-
lian ones.
DISCUSSION
Three hypotheses regarding the oviposi-
tional technique of Stylogaster were sug-
gested in the Introduction: (A) shooting the
eggs at the host from above with a flipping
movement of the abdomen; (B) stabbing
them into the host after a darting approach;
and (C) the more conventional method of
inserting an egg after seizing the victim.
The Results section reviewed the available
circumstantial evidence on the morphology
of the female postabdomen, the eggs, and
their placement in the host, and introduced
new evidence regarding the morphology of
the internal female reproductive tract. From
this evidence the following inferences can
be derived.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1. Stylogaster does not seize its host for
Oviposition. Otherwise the egg placement
would be expected to be more precise, as is
the case in other parasitic flies that seize
their hosts, such as Zodion obliquefascia-
tum Macquart (Conopidae, Howell 1967).
Oviposition on the head and wings would
presumably be avoided in favor of loca-
tions, such as the abdomen, where the cu-
ticle is thinner and the larva is likely to gain
access to the body tissue. Moreover, grasp-
ing or seizing of the host has never been
observed in Stylogaster. Therefore, the ev-
idence does not support hypothesis C.
2. The penetration of the host cuticle
must be achieved entirely by the tip of the
egg itself. The terminalia of Stylogaster fe-
males lack any specialized piercing struc-
tures. Instead, the eggs show conspicuous
adaptations for piercing, such as a strongly
sclerotized and sharply pointed tip.
3. For stabbing oviposition, a fertilized
egg would need to be pre-arranged and se-
cured in such a manner, that its pointed tip
protrudes from the female postabdomen.
This is indeed the case. In several Stylo-
gaster specimens, an egg was found in the
appropriate position, with the anterior egg
pole firmly lodged in the correspondingly
shaped terminal chamber, and the posterior,
pointed tip protruding from the vulva. It is
hard to envision how this position can be
achieved without the egg entirely slipping
out of the vulva. Apparently the inward
curved endlobes of the lateral appendages
of sternum 8 take part in securing the egg.
4. On its way from the oviduct to the
terminal chamber, the egg has to be fertil-
ized. The anteroventral evagination of Sty-
logaster is similarly positioned and perhaps
homologous to the ventral receptacle of
other acalyptrate Schizophora (Sturtevant
1925, 1926, Kotrba 1993) and the fertiliza-
tion chamber of the Calyptrates (Leopold et
al. 1978). It is conceivable that it likewise
receives the anterior egg pole sometime
during oviposition, possibly functioning as
the fertilization site. The conspicuous mus-
cle band that extends anteriorly from the
VOLUME 99, NUMBER 4
anteroventral evagination obviously pulls
this organ into an anterior position while
the vagina is extended. If the anteroventral
evagination receives the egg in this position
and the muscle band is relaxed subsequent-
ly, the egg will be propelled posteriorly by
the elastic shortening of the extended va-
gina. Such a mechanism could considerably
speed up the processes of fertilization and
transportation of the egg.
5. If the egg is already positioned in the
terminal chamber while the female is hov-
ering with the ovipositor extended posteri-
orly (Fig. 1), then the frequently observed
flicking movement of the postabdomen
could be correlated with accelerating and/or
launching the egg. The shooting hypothesis
(A) matches the suppositions of some eye-
witnesses and would account best for the
random egg placement. The flicking of the
abdomen is, however, unlikely to provide
the necessary kinetic energy for the pene-
tration of the host cuticle. So far, there are
no clues to the presence of any other mech-
anism to accelerate the eggs.
6. In the case of stabbing oviposition
(hypothesis B) the required energy could be
derived from the stabbing impact, which
might indeed be high due to the velocity of
the approaching flight, described as “‘dart-
ing’? or ‘“‘pouncing’’ by eye-witnesses.
Upon impact, the protruding posterior egg
pole is thrust into the host cuticle while the
anterior egg pole is prevented from giving
way anteriorly by the sclerotized roof of the
terminal chamber. After penetrating the host
cuticle, the egg is locked by its recurrent
barbs and pulled free of the oviscapt as the
Stylogaster female flies away.
While this manuscript was being re-
viewed, Elke Buschbeck (personal com-
munication) told me of an observation from
Costa Rica that corroborates the latter sce-
nario. She observed Stylogaster females
hovering about 30 cm above the ground at
the front of an army ant raid. One Stylo-
gaster female darted down upon a cock-
roach, which was quickly running from one
leaf to another for shelter. After an impact
621
of a fraction of a second the female flew
away, and a light-colored object, presum-
ably an egg, was visible sticking in the tho-
rax of the cockroach.
CONCLUSIONS
Lacking detailed observations, the ques-
tion of whether Stylogaster oviposits by
shooting (hypothesis A) or stabbing (hy-
pothesis B) can only be addressed through
evaluation of circumstantial evidence. The
present study favors the stabbing hypothesis
suggested by Smith (1967, 1979). Stabbing
is technically easier to conceive than shoot-
ing, and agrees well with the morphological
evidence and some of the behavioral obser-
vations. However, even considering the
high speed of a stabbing encounter, hypoth-
esis B does not account as well for the poor
egg placement on the host.
For a definitive answer, more detailed ob-
servations of oviposition in Stylogaster will
be essential. This requires field work in
South America or Africa, where Stylogaster
can be seen ovipositing at the front of raid-
ing army ant swarms. Alternatively, living
Stylogaster females could be collected and
provided with appropriate hosts, such as
cockroach larvae or silvicolous dung-breed-
ing Muscids, in a free-flying arena. In fu-
ture investigations of the female postabdo-
men special attention should be given to
structures that are apt to store and instantly
release energy, such as parts of resilin or a
notch and hook release mechanism.
ACKNOWLEDGMENTS
This study was supported by a postdoc-
toral fellowship from the Smithsonian In-
stitution, Washington D.C. I thank Margaret
Schneider for sending 5 Stylogaster frauci
females in alcohol. The photo of a hovering
Stylogaster female in Costa Rica (Fig. 1)
was contributed by Riley Nelson. Wayne
Mathis supervised and most graciously sup-
ported my work at the NMNH. I thank him
as well as Alessandra Baptista, Steven Lin-
gafelter, Riley Nelson, Kenneth Smith, Bri-
an Stuckenberg, Norman Woodley, and two
622
anonymous referees for critical comments
on the manuscript.
LITERATURE CITED
Bates, H. W. 1863. The Naturalist on the River Am-
azon. First edition, Volume 2. Murray, London. vi
+ 423 pp.
Carpenter |G: D> HS S153 Dr Gab sie Carpenters
observations on Dorylus nigricans Mlig., in Dam-
ba and Bugalla Islands. Proceedings of the Royal
Entomological Society of London 1914: evii-cxi.
de Meijere, J. C. H. 1904. Beitrage zur Kenntnis der
Biologie und der systematischen Verwandtschaft
der Conopiden. Tijdschrift voor Entomologie 46:
144-225 + plates.
Hardy, D. E. 1987. Pipunculidae, pp. 745-748. In
McALpine, J. E,ed., Manual of Nearctic Diptera,
Volume 2. Agriculture Canada, Research Branch
(Monograph; No. 28), Ottawa.
Hinton, H. E. 1981. Biology of insect eggs. 3 Volums.
Pergamon Press, Oxford. 1125 pp.
Howell, J. E 1967. Biology of Zodion obliquefascia-
tum (Macq.) (Diptera: Conopidae). Bulletin of the
Washington Agricultural Experiment Station 51:
33 pp.
Kotrba, M. 1993. Das Reproduktionssystem von Cyr-
todiopsis whitei Curran (Diopsidae, Diptera) unter
besonderer Beriicksichtigung der inneren wei-
blichen Geschlechtsorgane. Bonner zoologische
Monographien 33: 115 pp.
Leopold, R. A., S. Meola, and M. E. Degrugillier.
1978. The egg fertilization site within the house
fly, Musca domestica L. (Diptera: Muscidae). In-
ternational Journal of Insect Morphology and Em-
bryology 7: 111-120.
Lindner, E. 1955. Ostafrikanische Omphralidae, The-
revidae und Conopidae (Dipt.). Jahresheft. Verein
fiir Vaterlandische Naturkunde in Wiirttemberg
110: 19-23.
Lopes, H. de Souza. 1937. Contribuigaéo ao conheci-
mento do genero < Stylogaster >> Macquart,
1835. Archivos. Institudo Biologica 3 (2): 257—
298)
1938. Sobre uma nova espécie do genero
Stylogaster Macquart, do Brasil. Memorias do In-
stituto Oswaldo Cruz 33 (3): 403-405 + 1 plate.
1971. Notes on some old Species of Stylo-
gaster, especially on paratypes of Aldrich’s spe-
cies (Diptera, Conopidae). Anais. Academia
Brasileira de Ciéncias 43 (3/4): 691-710.
, and L. Monteiro. 1959. Sdbre algumas es-
pécies brasileiras de Stylogaster Macq., com a
descrigao de quatro espécies novas. Studia Ento-
mologica 2 (1—4): 1-24.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Rettenmeyer, C. W. 1961. Observations on the biol-
ogy and taxonomy of flies found over swarm raids
of army ants (Diptera: Tachinidae, Conopidae).
The University of Kansas Science Bulletin 42 (8):
993-1066.
Smith, K. G. V. 1966. The larva of Thecophoa occi-
densis, with comments upon the biology of Con-
opidae (Diptera). Journal of Zoology, London
149: 263-276.
. 1967. The biology and taxonomy of the ge-
nus Stylogaster Macquart, 1835 (Diptera: Conop-
idae, Stylogasterinae) in the ethiopian and mala-
gasy regions. Transactions of the Royal Entomo-
logical Society of London 119 (2): 47-69.
. 1969. Further data on the oviposition by the
genus Stylogaster Macquart (Diptera: Conopidae,
Stylogasterinae) upon adult calyptrate Diptera as-
sociated with ants and animal dung. Proceedings
of the Royal Entomological Society of London
(A) 44 (1-3): 35-37.
1979. The genus Stylogaster (Diptera: Co-
nopidae: Stylogasterinae) in the australian region.
Australian Journal of Zoology 27: 303-310.
, and G. R. Cunningham-van Someren. 1985.
The larva of Stylogaster varifrons Malloch (Dipt.,
Stylogasteridae). Entomologist’s Monthly Maga-
zine 121: 81-85.
, and B. V. Peterson. 1987. Conopidae, pp.
749-756. In McALpine, J. E, ed., Manual of Ne-
arctic Diptera, Volume 2. Agriculture Canada, Re-
search Branch (Monograph; No. 28), Ottawa.
Steyskal, G. C. 1987. Pyrgotidae, pp. 813-816. In
McALpine, J. FE, ed., Manual of Nearctic Diptera,
Volume 2. Agriculture Canada, Research Branch
(Monograph; No. 28), Ottawa.
Streng, R. 1976. Die Haarbalgmilbe des Menschen:
Demodex folliculorum. Mikrokosmos 7: 202-205.
Stuckenberg, B. R. 1963. A study on the biology of
the genus Stylogaster, with the description of a
new species from Madagascar (Diptera, Conopi-
dae). Revue de Zoologie et de Botanique Afri-
caines 28: 251-275.
Sturtevant, A. H. 1925. The seminal receptacles and
accessory glands of the Diptera, with special ref-
erence to the Acalypterae. Journal of the New
York Entomological Society 33: 195-215.
. 1926. The seminal receptacles and accessory
glands of the Diptera, with special reference to the
Acalyptratae. Journal of the New York Entomo-
logical Society 34: 1-21 + plates 1-3.
Townsend, C. H. T. 1897. Contributions from the New
Mexico Biological Station—No. 2. On a collec-
tion of Diptera from the lowlands of the Rio Nau-
tla, in the State of Vera Cruz. 1. Annals and Mag-
azine of Natural History (6) 19: 16-34.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 623-627
EMPOASCA (KYBOS) LUDA DAVIDSON AND DELONG: DISTRIBUTION AND
HABITS OF AN IMMIGRANT BIRCH-FEEDING LEAFHOPPER IN NORTH
AMERICA (HOMOPTERA: CICADELLIDAE)
A. G. WHEELER, JR.
Department of Entomology, Clemson University, Clemson, SC 29634, U.S.A.
Abstract.—The typhlocybine leafhopper Empoasca (Kybos) luda Davidson and De-
Long, though first described from the United States, is believed to be a native European
species that has been accidentally introduced into North America. New state records are
given for Maryland, New York, and Pennsylvania. Overwintering occurs in the egg stage,
and first-generation adults appear from late May through June; a second generation is
produced in mid- to late summer. Nymphs feed on lower leaf surfaces of European white
birch (Betula pendula Roth) in landscape plantings, causing chlorosis on the upper sur-
faces. Taxonomic characters are provided that allow this minor pest to be distinguished
from a generally more serious pest of ornamental birches, the often co-occurring potato
leafhopper, E. fabae (Harris).
Key Words:
Empoasca (Kybos) luda Davidson and
DeLong is a West European typhlocybine
leafhopper considered to have been acci-
dentally introduced into the Nearctic Re-
gion (Hamilton 1972, 1983, Hamilton and
Langor 1987). First collected in North
America in 1917 (Davidson and DeLong
1938), this birch feeder has remained little
known. Here, I review its taxonomic his-
tory, give new U.S. records, and provide
notes on its seasonality and habits on or-
namental birches. Characters are also pro-
vided that facilitate the recognition of E.
luda on birch trees.
TAXONOMIC HISTORY
Though regarded as adventive in the
New World, E. luda was first described
from North America; Davidson and De-
Long (1938) based their description on sin-
gle specimens from Illinois and Ohio. In
Europe, Wagner (1955) described the new
species E. betulicola from Germany, noting
Insecta, Cicadellidae, leafhoppers, distribution, birches, feeding habits
that it superficially resembles the Palearctic
E. smaragdula (Fallén). Hamilton (1983),
recognizing that the birch-feeding species
described by Wagner is conspecific with
that described by Davidson and DeLong,
proposed E. betulicola as a synonym of E.
luda. Ross (1963) incorrectly synonymized
E. luda under E. smaragdula (P. W. Oman,
pers. comm. 1989), which specializes on al-
ders (Alnus spp.) (e.g., Nuorteva 1952, Le
Quesne 1960, Claridge et al. 1968, Claridge
and Wilson 1976, Dworakowska 1976, Os-
siannilsson 1981).
Empoasca luda has been confused with
E. smaragdula in both Europe and North
America. Apparently the only valid Nearc-
tic record of the latter species is that of
Hamilton (1983) from Vancouver, B.C. The
records of Poos and Wheeler (1943), who
referred to E. smaragdula as widely distrib-
uted in Canada and occurring throughout
the United States, are based on misidenti-
fications (see Hamilton 1983). Their listing
624
of alder, birch, crabapple, linden, poplar,
and willow as host plants suggests that sev-
eral Empoasca species, including E. luda,
are contained in their concept of E. sma-
ragdula. Based on Hamilton (1983, 1985),
Beirne’s (1956) record of E. smaragdula
from Ontario is also misidentified and re-
fers to E. luda.
Kybos and other subgenera of Empoasca
are sometimes given generic status (e.g.,
Dlabola 1958, Le Quesne 1960). Here,
however, I follow Ossiannilsson (1981),
Hamilton (1983), and others who consider
Kybos a subgenus of Empoasca.
NORTH AMERICAN DISTRIBUTION
In addition to Illinois and Ohio (David-
son and DeLong 1938), E. luda is known
from Connecticut, Minnesota, Ontario, and
Virginia (Metcalf 1968). Subsequent to the
Metcalf catalogue, it has been reported
from British Columbia, Ontario (Hamilton
1972, 1983), and Newfoundland (Hamilton
and Langor 1987). The following new state
records are based on my collections from
ornamental Betula pendula Roth. Voucher
specimens have been deposited in the col-
lections of Cornell University, Ithaca, N.Y.;
National Museum of Natural History,
Washington, D.C. (USNM); and the Penn-
sylvania Department of Agriculture, Harris-
burg.
MARYLAND: Allegany Co., Frostburg
State University, Frostburg, 28 July 1989,
9 May 1990, 3 June 1990; Frostburg, 23
May 1993 (nymphs only); Zihlman, 28 July
1989. NEW YORK: Tompkins Co., Ithaca,
29 Aug. 1989. PENNSYLVANIA: Berks
Co., Kutztown University, Kutztown, 18
Aug. 1989; Huntingdon Co., Greenwood
Furnace, 28 June 1990; Lehigh Co., Trex-
lertown, 8 Aug. 1989; Luzerne Co., Wilkes-
Barre, 14 Aug. 1989; Northampton Co.,
Cherryville, 8 Aug. 1989.
Empoasca luda has been recorded from
northern Virginia (Arlington) by Wheeler
(1942). I have collected it on B. pendula in
southwestern Virginia (Montgomery Co.,
Virginia Tech campus, Blacksburg, 2 July
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1995), which is the southernmost record of
this adventive leafhopper in North America.
SEASONAL HISTORY, HOSTS, AND HABITS
Notes on seasonality are based on collec-
tions and observations of small numbers of
individuals (often 10 or fewer) made peri-
odically from European white (or silver)
birch (B. pendula), the white-barked birch
most commonly planted as an ornamental
in the United States. In the horticultural
trade, this species, as well as the European
B. pubescens Ehrh., has often been called
B. alba L. (Santamour and McArdle 1989).
At Trexlertown, Pa., E. Juda was found on
a cut-leaved, pendulous birch, probably B.
pendula ‘laciniata’.
The collection of second-and third-instar
nymphs in early May in western Maryland
suggests that E. Juda overwinters in the egg
stage, as does E. smaragdula and apparent-
ly other members of the subgenus Kybos
(Nuorteva 1952, Ossiannilsson 1981).
Fourth instars were observed in late May,
and in early June about equal numbers of
late-instar nymphs and adults were present
at Frostburg, Md. In Pennsylvania, adults
and late instars of a first generation were
found in late June. The collection of
nymphs in early to mid-August in Pennsyl-
vania and in late August in New York sug-
gests the occurrence of two generations on
ornamental birches. At Arlington, Va.,
adults have been collected as late as 8 Oct.
(USNM collection). This leafhopper is also
thought to be bivoltine in Europe (Dwora-
kowska 1976).
Nymphs occur mainly on the abaxial sur-
face of birch leaves, their feeding on me-
sophyll producing chlorosis on the adaxial
surface (Fig. 1). In early season, and when
relatively small numbers of E. Juda are
present, the injury consists of light stip-
pling. Only in western Maryland in 1989
was chlorosis conspicuous on the foliage of
European white birch.
RECOGNITION FEATURES
Wild and cultivated birches, including B.
pendula, serve as hosts of numerous leaf-
VOLUME 99, NUMBER 4
lene, Ae
caused by the mesophyll-feeding leafhopper Empoasca
(Kybos) luda.
Chlorosis on foliage of Betula pendula
hoppers in North America (Varty 1967,
Hamilton 1985), but E. luda is larger (4.2—
4.8 mm) than most other birch-feeding ty-
phlocybines, and it is the only cicadellid
species likely to occur on European white
birch that has greenish and brown- or black-
tinged forewings. Hamilton (1985) noted
that adults are green or orange with black
stripes, but all specimens I have seen are
green or yellowish green with dark mark-
ings. Empoasca fabae (Harris), the potato
leafhopper, frequently co-occurs with E.
luda, but it is more delicate and smaller
(about 3.0—3.5 mm), generally pale green
without dark markings, and has a bluntly
angled (rather than broadly rounded) ver-
tex. Although E. Juda has a distinctive hab-
itus among leafhoppers associated with or-
namental white birch, it cannot be separated
reliably from other typhocybines, especially
other members of the diverse genus Em-
poasca, without examining the male geni-
625
talia (Figs. 2—9). Male genitalic characters
allowing E. luda (as E. betulicola) to be
distinguished from the presumed closely re-
lated E. smaragdula are discussed and il-
lustrated in LeQuesne (1961), Dworakow-
ska (1976), and Ossiannilsson (1981).
DISCUSSION
Empoasca luda is one of the relatively
few Old World insects now established in
the New World that was first described
from North America; the mullein thrips,
Haplothrips verbasci (Osborn), is believed
to represent a similar example (Stannard
1968). The birch specialist E. Juda is one
of numerous Palearctic leafhoppers sug-
gested to have been accidentally introduced
into North America with shipments of nurs-
ery stock (Hamilton 1983, Hamilton and
Langor 1987). The adventive status of E.
luda in the Nearctic region is supported by
my failure to collect this species in north-
eastern North America on native birches
such as B. lenta L., B. papyrifera Marsh.,
or B. populifolia Marsh. Within the Holarc-
tic subgenus Kybos, an association with Al-
nus and Betula (Betulaceae) is considered
secondary, the ancestral hosts belonging to
either Populus or Salix (Salicaceae) (Ross
1963).
Empoasca luda has not been reported
previously as a pest of ornamental birches.
As a mesophyll feeder, it causes chlorosis
on the foliage of European white birch.
Chlorosis is typically restricted to relatively
few branches, although in 1989 injury was
widespread on three trees in a Maryland
landscape planting. Horticulturists, plant in-
spectors, and pest-management specialists,
therefore, should be aware of this leafhop-
per’s potential for causing aesthetic injury
to European white birch. They should not,
however, mistake E. luda for E. fabae, a
mesophyll and occasional phloem feeder
(e.g., Backus 1989) that can cause more se-
rious problems on ornamental birches—
stunting, distortion, and swelling of twigs—
in nurseries and landscape plantings (Tash-
iro 1973, Valley and Blosser 1986, Johnson
626
Figs. 2-9.
1
!
ved
!
1
'
|
|
'
1
!
!
!
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
I SS A os
Empoasca (Kybos) luda, male terminalia. 2, Left anal collar appendage from left. 3, Left pygofer
appendage from outside. 4, Aedeagus from left. 5, Aedeagus in ventral aspect. 6, Aedeagus in ventral aspect.
7, 2nd abdominal sternum from below. 8, 2nd abdominal tergum from above. 9, 3rd abdominal tergum from
below. Scale = 0.1 mm (From Ossiannilsson 1981:436).
and Lyon 1988). I agree with Raupp (1990),
who emphasized that managing arthropod
pests of woody ornamentals depends on ac-
curate identification of both the host plants
and their arthropod associates, pests as well
as beneficial and neutral species. Empoasca
luda should be considered an occasional
minor pest of European white birch.
ACKNOWLEDGMENTS
I gratefully acknowledge E. R. Hoebeke
(Cornell University) and P- W. Oman (Or-
egon State University) for confirming the
identification of E. luda; T. E. Wolf (Penn-
sylvania Department of Agriculture) for
photographing leafhopper injury; J. EK Stim-
mel (PDA) for preparing Fig. 1; T. J. Henry
(Systematic Entomology Laboratory
USDA) for checking the USNM collection
for specimens of FE. Juda; P. H. Adler and
J. C. Morse (Clemson University) for re-
viewing an early draft of the manuscript;
and the publisher, E. J. Brill, Leiden, The
Netherlands, for granting permission to re-
VOLUME 99, NUMBER 4
produce figs. 1394-1401 from Ossiannils-
son’s 1981 paper. This is Technical Contri-
bution No. 4256 of the South Carolina Ag-
ricultural Experiment Station, Clemson
University.
LITERATURE CITED
Backus, E. A. 1989. Host acceptance and feeding be-
havior, pp. 10-17. Jn Armbrust, E. J. and W. O.
Lamp, eds., Proceedings of a Symposium: History
and Perspectives of Potato Leafhopper (Homop-
tera: Cicadellidae) Research. Entomological So-
ciety of America, Lanham, Md.
Beirne, B. P. 1956. Leafhoppers (Homoptera: Cica-
dellidae) of Canada and Alaska. Canadian Ento-
mologist 88 (supplement 2): 5-177.
Claridge, M. F, J. M. Edington, and D. M. Murphy.
1968. The distribution of some Hemiptera in the
birch-dominated woodlands of northern Scotland.
Entomologist 101: 253-263.
Claridge, M. EK and M. R. Wilson. 1976. Diversity and
distribution patterns of some mesophyll-feeding
leafhoppers of temperate woodland canopy. Eco-
logical Entomology 1: 231-250.
Davidson, R. H. and D. M. DeLong. 1938. Studies on
the genus Empoasca (Homoptera, Cicadellidae).
Part V. Twelve new species of Empoasca from the
United States. Ohio Journal of Science 38: 90-96.
Dlabola, J. 1958. A reclassification of Palaearctic Ty-
phlocybinae (Homopt., Auchenorrh.). Acta Socie-
tatis Entomologicae Cechosloveniae 55: 44—57.
Dworakowska, I. 1976. Kybos Fieb., subgenus of Em-
poasca Walsh (Auchenorryncha, Cicadellidae,
Typhlocybinae) in Palaearctic. Acta Zoologica
Cracoviensia 21: 387—463.
Hamilton, K. G. A. 1972. The leafhopper genus Em-
poasca subgenus Kybos in the southern interior of
British Columbia. Journal of the Entomological
Society of British Columbia 69: 58-67.
Hamilton, K. G. A. 1983. Introduced and native leaf-
hoppers common to the Old and New Worlds
(Rhynchota: Homoptera: Cicadellidae). Canadian
Entomologist 115: 473-511.
Hamilton, K. G. A. 1985. Leafhoppers of ornamental
and fruit trees in Canada. Agriculture Canada
Publication 1779/E. 71 pp.
Hamilton, K. G. A. and D. W. Langor. 1987. Leaf-
hopper fauna of Newfoundland and Cape Breton
Islands (Rhynchota: Homoptera: Cicadellidae).
Canadian Entomologist 119: 663-695.
Johnson, W. T. and H. H. Lyon. 1988. Insects that
Feed on Trees and Shrubs, 2nd ed. Cornell Uni-
versity Press, Ithaca, N.Y. 556 pp.
627
LeQuesne, W. J. 1961. An examination of the British
species of Empoasca sensu lato (Hem., Cicadel-
lidae), including some additions to the British list.
Entomologist’s Monthly Magazine 96: 233-239.
Metcalf, Z. P. 1968. General Catalogue of the Ho-
moptera. Fascicle VI Cicadelloidea, Part 17 Ci-
cadellidae. Agricultural Research Service, U.S.
Department of Agriculture, Washington, D.C.
1513 pp.
Nuorteva, P. 1952. Die Nahrungspflanzenwahl der In-
sekten im Lichte von Untersuchungen an Zikaden.
Annales Academiae Scientiarum Fennicae IV
Biologica 19(A): 1—90.
Ossiannilsson, F 1981. The Auchenorrhyncha (Ho-
moptera) of Fennoscandia and Denmark. Part 2:
The families Cicadidae, Cercopidae, Membraci-
dae, and Cicadellidae (excl. Deltocephalinae).
Fauna Entomologica Scandinavica 7(2): 223-593.
Poos, EK W. and N. H. Wheeler. 1943. Studies on host
plants of the leafhoppers of the genus Empoasca.
U.S. Department of Agriculture Technical Bulletin
850. 51 pp.
Raupp, M. J. 1990. Recognizing the larvae of key
pests and beneficials found on woody landscape
plants. Journal of Arboriculture 16(3): 49-54.
Ross H. H. 1963. An evolutionary outline of the leaf-
hopper genus Empoasca subgenus Kybos, with a
key to the Nearctic fauna (Hemiptera, Cicadelli-
dae). Annals of the Entomological Society of
America 56: 202-223.
Santamour, E S., Jr. and A. J. McArdle. 1989. Check-
lists of cultivars in Betula (birch). Journal of Ar-
boriculture 15: 170-176.
Stannard, L. J. 1968. The thrips, or Thysanoptera, of
Illinois. Illinois Natural History Survey Bulletin
29(4): 215-552.
Tashiro, H. 1973. Evaluation of soil applied systemic
insecticides on insects of white birch in nurseries.
Search Agriculture. Entomology (Geneva, New
York) 3(9): 1=11.
Valley, K. and W. E. Blosser, Jr. 1986. Potato leaf-
hopper, Empoasca fabae (Harris), as a pest of or-
namentals (Homoptera: Cicadellidae). Regulatory
Horticulture 12(2): 11-12.
Varty, I. W. 1967. Leafhoppers of the subfamily Typh-
locybinae from birches. Canadian Entomologist
99: 170-180.
Wagner, W. 1955. Neue mitteleuropaéische Zikaden
und Blattfl6he (Homoptera). Entomologische Mit-
teilungen. Zoologischen Staatsinstitut und Zoolo-
gischen Museum (Hamburg) No. 6: 1-34.
Wheeler, N. H. 1942. Trap-light studies on leafhop-
pers of the genus Empoasca (Homoptera: Cica-
dellidae), 1932-1941. Proceedings of the Ento-
mological Society of Washington 44: 69-72.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 628-640
THE GENUS PLESKEOLA STACKELBERG (DIPTERA: SYRPHIDAE), A
JUNIOR SYNONYM OF PARHELOPHILUS GIRSCHNER
E CHRISTIAN THOMPSON
Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart-
ment of Agriculture, c/o National Museum of Natural History, MRC-168, Washington,
DC 20560, U.S.A.
Abstract.—The genus Pleskeola (n. syn.) is synonymized with Parhelophilus. The spe-
cies (porcus and sibiricus, n. comb.) associated with the name Pleskeola are redescribed.
A checklist and key to species of Parhelophilus is presented. Four new synonyms and
one new combination are reported (Helophilus anniae Brimley 1923 = flavifacies Bigot
1883; Parhelophilus obscurior Violovitsh 1960 and Helophilus citricornis Shiraki 1968
= kurentzovi Violovitsh 1960; Parhelophilus currani Fluke 1953 = laetus Loew 1863;
Helophilus pilosus Hunter 1897 is transferred to Lejops (Lejops)).
Key Words: key, holarctic, bogs
This is a tale of two rare flower flies,
which are known from only a few speci-
mens collected in the boreal forest. One
flower fly was the basis of a genus-group
name and is currently known only from
three specimens collected in Finland and
Siberia. The other flower fly is known from
more specimens, about 100 in all, and ap-
pears to be restricted to bogs in North
America. As part of a revision of the higher
classification of flower flies, I have sought
to re-examine the types of all genus-group
names and to place those types into my
classification. This eventually brought me
to Pleskeola sibirica Stackelberg, a genus
and species-group name proposed as new
by Stackelberg for a unique male collected
in central Siberia. On seeing the second
known specimen of Pleskeola, I immediate-
ly recognized it as being close, if not the
same as a rare bog fly (Parhelophilus por-
cus (Walker)) that I had once collected in
northern New Hampshire. While both por-
cus and sibiricus are unusual and share dis-
tinctive characters, the two species other-
wise agree with the characteristics of Par-
helophilus. Why Pleskeola is here reduced
to a junior synonym of Parhelophilus is
outlined below. What remains of interest is
why the species upon which the genus-
group name is based is so rare in collec-
tions. Is sibirica a bog species, like its sister
(porcus), and merely rare because northern
bogs are poorly collected in Russia? Per-
haps this treatment of these two species will
encourage field people to more closely ex-
amine bogs and look for these unusual flies.
The terminology and characters used
generally follow those used in the Manual
of Nearctic Diptera and all my standards are
defined and illustrated in a comprehensive
glossary (Thompson 1997). The format of
the checklist follows the standards of the
Biosystematic Database of World Diptera
(see Thompson 1997).
Genus Parhelophilus Girschner
Helophilus subg. Parhelophilus Girschner
1897: 604. Type species, Syrphus frute-
torum Fabricius by subsequent designa-
VOLUME 99, NUMBER 4
HEATHER
Quan
Fig. 1. Parhelophilus sibiricus, male, habitus, dorsal view.
tion of Curran and Fluke 1926: 230).
Curran and Fluke 1926: 230 (revision,
Nearctic spp.); Goffe 1944: 129 (nomen-
clature), 1945: 276 (type species); Hull
1949: 387 (description).
Pleskeola Stackelberg 1924: 25. Type spe-
cies, Pleskeola sibirica Stackelberg by
original designation. Sack 1931: 276 (de-
scription); Hull 1949: 402 (diagnosis);
Stackelberg 1970: 80, 1988: 121 (refer-
ence in key); Goot 1981: 211 (reference
in key); Violovitsh 1983: 114, 129 (ref-
erence in key). New synonym.
Head: Face pale, yellow to white, nar-
row, slightly longer than broad, occupying
about % of head width, straight to concave
beneath antennae, without distinct medial
tubercle, pollinose and pilose laterally, pol-
linose and bare medially; frontal promi-
nence low, at dorsal % of head; vertex
black, pollinose, pilose, rectangular, slightly
longer than wide; ocellar triangle variable,
from broader than long to longer than
broad; eye bare or pilose; always dichoptic
in males; antenna short, less than % as long
as face; basoflagellomere oval; arista bare,
slightly longer than antenna.
Thorax: Broader than long, long pilose,
usually with longitudinal pale pollinose vit-
tae on mesonotum (absent or greatly re-
duced in obsoletus); meso-katepisternum
continuously pilose from ventral to dorsal
margins; metathoracic spiracle small,
slightly smaller than basoflagellomere;
meso-anepimeron bare posteriorly; katepi-
meron bare; meron bare; metasternum usu-
ally pilose (bare in divisus); plumula elon-
gate, unbranched; subscutellar fringe ab-
sent. Legs: Hind femur slightly swollen;
hind tibia slightly arcuate, with basoventral
carina on basal % or less, without apical
calcar. Wing: Microtrichose; cell R1 open;
630
cell R4+5 with short petiole, with petiole
about as long as crossvein h; stigmatic
crossvein present.
Abdomen: Suboval, with paired lateral
light colored pollinose maculae on dorsum.
Girschner (1897), on the basis of a study
of the calypter of higher Diptera, divided
the genus Helophilus into subgenera. He
defined most of these subgenera with char-
acters, but Parhelophilus was defined by
exclusion (that is, those species which did
not fit the other defined subgenera). At first,
most workers ignored Girschner’s action as
they could not understand the characters on
which the group was based and, hence,
treated Parhelophilus as a synonym of He-
lophilus. As Verrall (1901: 524-525) noted,
Parhelophilus, as construed by Girschner,
was a heterogeneous group (two species be-
longed to Parhelophilus of current authors,
one to Lejops Rondani). Curran and Fluke
(1926) designated a type species, recog-
nized Parhelophilus as a distinct group, and
treated the group as a genus. North Amer-
ican workers have followed Curran and
Fluke as have some Palaearctic workers.
The characters that separate Parhelophilus
from Helophilus are: 1) presence of stig-
matic crossvein; 2) entirely pollinose face,
without a shiny medial vitta; and 3) short,
suboval and compact abdomen. Parhelo-
philus differs from Lejops in the 4) lack of
an apical calcar on the hind tibia as well as
the suboval abdomen. Pleskeola shares all
these characteristics with Parhelophilus.
Stackelberg (1924) erected his genus
Pleskeola on the basis of the pilose eyes,
an unusual character state among species
related to Helophilus (he also noted the
slightly sinuate vein R4+5 and in contrast
to Mallota, the dichoptic males). Stackel-
berg’s original description was brief and in
Latin. Because the species on which the ge-
nus was based was rare, most subsequent
workers merely repeated the original de-
scription and/or left Pleskeola as unplaced
or a genus of dubious status.
Eye pilosity is a character of variable im-
portance among flower flies. In most gen-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
era, the eyes of all included species are ei-
ther bare or pilose. Within some genera,
however, the character is variable, with
some species being bare eyed and others
pilose (e.g., Syrphus). In some species, the
male eyes are pilose and the females’ are
bare (e.g., Cheilosia latifrons (Zetterstedt)).
The eye pilosity of sibirica is irrelevant as
a genus-group character as other characters
reveal a sister-group relationship to one
species of Parhelophilus and common re-
lationships with all species of Parhelophi-
lus. Parhelophilus porcus and_ sibiricus
have the Ist and 2nd sterna (figs. 11—12)
fused and a distinct dark pollinose body
pattern (figs. 1-2), both characters found no
where else among the helophiline flower
flies. These two species share a character-
istic basoventral process on the hind femur
(figs. 14-15) in males with fruterorum and
divisus. As noted above, sibirica shares all
the characters of the genus Parhelophilus.
Hence, Pleskeola is here considered a syn-
onym of Parhelophilus (new synonym).
Parhelophilus porcus (Walker)
(Figs. 2,°6, 7, 10) MiSs ie
Eumerus porcus Walker 1849: 554 @ [error
for ¢] Ontario, Hudson’s Bay, Albany
River, St. Martin’s Falls (LT ¢ BMNH
here designated). Osten Sacken 1858: 48
(catalog citation), 1875: 61 (catalog ci-
tation, ?=Eristalis).
Helophilus porcus: Osten Sacken 1878:
134, 250 (catalog citation, note on types);
Williston 1887: 197 (description); Al-
drich 1905: 394 (catalog citation); Os-
burn 1907: 3 (British Columbia), 1908:
10 (British Columbia); Graenicher 1910:
41 (Wisconsin); Fluke 1922: 245 (Wis-
consin); Johnson 1923: 10 (Maine),
1925: 174 (New England); Wirth et al.
1965: 619 (catalog citation).
Tubifera porcus: Kertész 1910: 259 (cata-
log citation).
Parhelophilus porcus: Curran and Fluke
1926: 232 (description, distribution, fig-
ures (head, abdomen)); Petch and Mal-
thais 1932: 51 (Quebec); Strickland
VOLUME 99, NUMBER 4 631
HEATHER
LALLIAN,
Figs. 2-10. Features. 2, Parhelophilus porcus, habitus, dorsal view. 3, P. sibiricus, male, head, frontal view.
4, P. sibirica, male, head, lateral view. 5, P. sibiricus, male genitalia, 9th tergum and associated parts, lateral
view. 6, P. porcus, male genitalia, 9th tergum and associated parts, lateral view. 7, P. porcus, male genitalia,
9th sternum and associated parts, lateral view. 8, P. sibiricus, male genitalia, 9th sternum and associated parts,
lateral view. 9, P. sibiricus, male genitalia, 9th tergum and associated parts, dorsal view. 10, P. porcus, male
genitalia, 9th tergum and associated parts, dorsal view.
632
1938: 203 (Alberta); Cole & Schlinger
1969: 326 (descriptive note, distribution
western North America); Miliczky and
Osgood 1979: 21 (Maine, flower (Vac-
cinum spp.)).
Head: Face yellowish white, white pol-
linose and pilose; gena black, shiny and
bare anteriorly, white pilose and grayish-
white pollinose posteriorly; front black,
gray pollinose, black pilose; frontal lunule
yellow; vertex black, black pollinose and
pilose except for some yellow pile on ocel-
lar triangle; occiput black, grayish-white
pollinose, black pilose dorsally except some
yellow pile posterior to ocellar triangle,
white pilose ventrally. Eye bare. Antenna
orange, black pilose.
Thorax: Black, mainly yellow pilose,
sparsely gray pollinose, with black polli-
nose vittate pattern on mesonotum (see Fig.
2); postpronotum yellow pilose; mesono-
tum with intermixed black pile; anepister-
num black pilose dorsoposteriorly; halter
yellow; calypter and plumula white; scutel-
lum black, dull black pollinose except
broadly shiny marginally, black pilose ex-
cept yellow pilose marginally. Wing micro-
trichose. Legs mainly black and black pi-
lose; femoral-tibial joints of fore and mid
legs broadly orange; fore and mid tarsi yel-
low, pale pilose; white pilose on coxae,
posteriorly on mid femur, anterobasally on
hind femur. Hind femur of male with ba-
soventral process (Fig. 15).
Abdomen: Black, mainly black pilose;
tergum | bluish-gray pollinose except black
pollinose macula apicolaterally, short black
pilose except white pilose laterally; tergum
2 with mediolateral small orange macula,
shiny apically, elsewhere black pollinose
except for medial white pollinose macula,
yellow pilose on pale areas and basolater-
ally; terga 3 & 4 black pollinose except
shiny apically and with medial grayish
white pollinose macula, yellow pilose ba-
solaterally; tergum 5 gray pollinose, white
pilose; genitalia white pilose, sparsely pol-
linose; sterna black, subshiny, very sparsely
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
pollinose, shiny medially, white pilose; 2nd
& 3rd sterna of male with apicomedial
patches of long curved bristles (Fig. 11).
Male genitalia (Figs. 6, 7 & 10): surstyle
with broad posteromedial cleft and without
a dorsomedial notch on dorsal lobe; supe-
rior lobe short, only % as long as hypan-
drium.
Type.—Eumerus porcus Walker, lecto-
type ¢ BMNH, London, labeled ‘‘Hudson’s
Bay [reverse] 47 14,”’ “‘Hudson’s Bay, pres.
by, G. Barnston, 44. 17,’ and with the ap-
propriate Austen labels. I accept this male
as a syntype as Walker usually sexed flower
flies by the condition of the eyes. Hence,
the dichoptic male of porcus would appear
to Walker as a female. Osten Sacken (1878)
reported two “well-preserved specimens”
in the British Museum, unfortunately only
one was found and it is now headless.
Flight period.—May-—July.
Distribution.—British Columbia to New
Brunswick, south to Wisconsin and Mary-
land. The southernmost record (Maryland)
is from the 1965 Diptera catalog (Wirth et
al., 1965: 619). I have not found its source.
The Pennsylvania records from “North
Mountain” are from Ricketts near Lake
Ganoga (Johnson 1914).
Biology.—Nothing is known of the im-
mature stages of porcus, but by phyloge-
netic inference these will be aquatic as all
eristaliine flower fly larval are of the rat-
tailed maggot type. The adults are collected
in association with bogs, where they have
been frequently collected on flowers (Fra-
garia, Houstonia, Ledum groenlandicum,
Potentilla, and Rubus). Most of the known
sites for porcus are the typical kettlehole
bogs (Bingham Pond, Wilson Mills), but
some (Laurel Lake site) are better described
as fens bordered by spruce, fir and hemlock
(see Johnson 1985 for details on the bogs
of northeastern USA). Unfortunately, for
most collected specimens, no site informa-
tion is available.
Material Examined.—ALBERTA: Banff,
4 July 1922, C. B. D. Garrett (1 2 Curran
& Fluke 1926: 232). MANITOBA: Cedar
VOLUME 99, NUMBER 4
Lake, July 1993, Brues (1 6 USNM); Gim-
li, 13 June 1923, A. J. Hunter (1 d - Curran
& Fluke 1926: 232). ONTARIO: Ottawa (1
6 USNM); Lake Abitibi, Low Bush, 3 July
1925, N. K. Bigelow (1 6 MCZ); Mc-
Diarmid (Curran & Fluke 1926: 232).
QUEBEC: Montreal, June 15 1906 (1 @
Curran & Fluke 1926: 232). CONNECTI-
CUT. Litchfield Co., Salibury, 5 km NW of
town center, bog at edge of Bingham Pond,
1,894 ft, 14-16 June 1983 (5 56 CTM), 20
June 1984 (1 6 CTM), 23 May 1985 (2 ¢
1 2 CTM), 4 June 1986 (1 ¢ 1 2 CTM),
all specimens collected in Malaise trap
among ericaceous shrubs at the edge of a
pond surrounded by bog mat with spruces.
MAINE. Oxford Co., Lincoln Plantation,
Maine highway 16, 6 miles south of Wil-
sons Mills, ““Wilson Mills Bog,”’ [a sphag-
num bog dominated by black spuce and
larch]: 4 June 1976 (1 do FEB), E D. Fee;
19 June 1982 on Ledum groenlandicum (2
6 FEE) EF Fee, F D. Fee (2 6 FEE); 24
June 1982 on Ledum groenlandicum (1 3
52 BEE), FD: Fee, F Fee (1 ¢ FEE): 6
June 1986 on Ledum groenlandicum, F. D.
Fee (1 6 FEE); 29 May 1987 on or about
Fragaria, F D. Fee (1 6 1 2 FEE); 3 June
1989 on Fragaria, FE D. Fee (1 ¢ FEE); 19
June 1982, C. T. Maier (1 6 CTM). Penob-
scot Co., Passadumkeag, 26 May 1977, E.
Miliczky (1d USNM). NEW YORK: Essex
Co., Lake Champlain, Corlaer Bay, June
1939 (1 6 1 2 USNM). NEW HAMP-
SHIRE. Coos Co.: Errol, 8 km SW junction
of New Hampshire highways 16 and 26, 19
June 1982, C. T. Maier (1 teneral 2 CTM);
Pittsburg, Rt. 3, Connecticut Lakes, 22—24
June 1972 B. J. & E C. Thompson (1 6
USNM); Pittsburg, Back Lake, 17 June
1982 on Fragaria (2 2° FEE), 22 June 1982
CNS 12S EEE); 18 June 1983 d Gal. °
FEE), 3 June 1986 on Fragaria (2 3 1
FEB), all collected by E D. Fee; Clarksville,
Hurlbert Swamp, 15 June 1984 E D. Fee (1
6 FEE); 2nd Connecticut Lake, Route 3
west of Dam, 3—5 June 1988 on Fragaria
(1 3d FEE) and on Ledum groenlandicum (1
6 1 2 FEB), all collected by ED: Fee;
633
Connecticut Lakes, Scott Bog, 8 June 1990
on Fragaria (1 3 FEE), 14 June 1990 on
Taraxacum (1 ¢ FEE), both collected by E
D. Fee. Sullivan Co.: Cornish Flat, 12 July
1971 A. G. Lavallee (1 6 12: USNM):
VERMONT, Windham Co., Laurel Lake,
near Jacksonville, 30 May 1977, on Hous-
tonia, H. D. Pratt (1 ¢6 1 2 USNM). PENN-
SYLVANIA. Centre Co.: Black Moshan-
non State Park, 15 May 1977 (1 2 FEE),
11 June 1977 (1 2 FEE), 16 June 1978 (1
3 2 2 FEE), 22 May 1979 on Taraxacum
(1 2 FEE), 13 June 1979 on Rubus (3 6 1
2 FEB), all collected by E D. Fee; Bear
Meadows Natural Area, 17 June 1987 on
Ranunculus (1 3 FEE), 26 June 1989 (1
FEE), all collected by E D. Fee. Tioga Co.:
Morris, 6 km Northwest of, State Game
Lands #268, 15 June 1979 in boggy sedge
meadow with some open water (2 6 FEE),
20 June 1979 (2 3 FEE), 4 June 1980 on
Rubus and Potentilla (3 36 3 2 FEE), 13
June 1980 on Rubus (2 5 2 2 FEE), 23
June 1980 on Rubus (1 ¢o FEE), 1 June
1981 on Rubus & Fragaria (3 2? FEE), 7
June 1981 on Rubus & Potentilla (4 3 1 9
FEE), all collected by F D. Fee: Euzeme
Co., North Mountain [=Ricketts, near Lake
Ganoga], June 6 ?? (1 d USNM), June 4
2? @*? MCZ), June s 1898 @xg° 22
USNM, 1 ¢ 1 2 MCZ) all collected by C.
W. Johnson.
Parhelophilus sibiricus (Stackelberg),
NEW COMBINATION
(Figs. 1, 3-5, 8, 9, 12, 14)
Pleskeola_ sibirica Stackelberg 1924: 25.
Type-locality: Russia, “Sibiria centralis,
Obj-Enisseijskij kanal. septen. et orien-
tem versus Enisseisk.”” HT ¢ ZISP. Sack
1931: 276 (description); Stackelberg
1970: 80, 1988: 121 (description, figures
(abdomen, wing); Goot 1981: 211 (ref-
erence in key, figures (abdomen, wing);
Violovitsh 1983: 129, 1986: 123 (Sibe-
ria); Peck 1988: 193 (cat. cit.)
Head (Figs. 3, 4): Face yellowish white,
white pollinose and pilose; gena black,
634 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 11-15. Features, ventrolateral views. 11-12, Abdomen, male. 11, Parhelophilus porcus. 12, P. sibiricus.
13-15, Hind leg, lateral view. 13, P. porcus, female. 14, P. sibiricus, male. 15, P. porcus, male.
VOLUME 99, NUMBER 4
shiny and bare anteriorly, yellow pilose and
gray pollinose posteriorly; front yellowish
white, white pollinose and pilose; frontal
lunule yellow; vertex black, black pollinose
and pilose except for some yellow pile on
ocellar triangle; occiput black, gray polli-
nose, black pilose dorsally except some yel-
low pile posterior to ocellar triangle, yellow
pilose ventrally. Eye brown pilose. Antenna
brown, black pilose.
Thorax: Black, mainly black pilose,
sparsely gray pollinose, with darker polli-
nose vittate pattern on mesonotum (see Fig.
1); postpronotum yellow pilose; mesono-
tum with intermixed yellow pile; katepi-
sternum yellow pilose; halter yellow; calyp-
ter and plumula white; scutellum black, dull
brown pollinose except shiny marginally,
black pilose. Wing microtrichose. Legs
black, mainly black pilose; yellow pilose on
coxae, posteriorly on mid femur, anteroba-
sally on hind femur. Hind femur of male
with basoventral process (Fig. 14).
Abdomen: Black, mainly black pilose;
tergum | black pollinose, short black pilose
except white pilose laterally; tergum 2 with
mediolateral small orange macula, shiny
apically, else where black pollinose except
for medial white pollinose macula, yellow
pilose on pale areas and basolaterally; terga
3 and 4 black pollinose except shiny api-
cally and with medial grayish white polli-
nose macula, yellow pilose basolaterally;
genitalia black pilose, dull pollinose; sterna
black, subshiny, very sparsely pollinose,
white pilose; sterna of male simple (Fig.
12). Male genitalia (Figs. 5, 7, and 9): sur-
stylus without posteromedial cleft, with
small medial notch on mesial side of dorsal
lobe; superior lobe elongate, longer than
hypandrium.
Distribution.—Russia (Karelia & Sibe-
ria).
Material examined.—RUSSIA. Karelia:
‘‘Paanajarvi [=Ozero Olanga, 66°8'N
30°0’E]; 833, R. Frey’’ (UZMH, 1 ¢). Si-
beria: ““23 May 1908, Tshugunov, Obj En-
issejskij Canal, NE of Enisseisk’’ [= Ob’ -
635
Yenisey rivers Canal, northeast of Yeni-
seysk](ZISP, Holotype ¢).
Stackelberg (1970: 80) listed Finland on
the basis of the Frey specimen. Hackman
(1980) did not list the species from Finland
because the Frey specimen, which was col-
lected in Finland, was collected at a locality
which is now in Russia! This specimen was
unknown to Sack (1931) and Violovitsh
(1983, 1986), who stated that it was known
only from the unique original male. Peck
(1988) gave Yakutia as additional locality.
Unfortunately, I have been unable to find
the specimen on which this record is based.
KEY TO THE SPECIES OF PARHELOPHILUS
ly Hem alee are Reka, Siete: ce aes. © Lh hie ree so. 15
INA Six, ANS coy aioe eas, EN Ee arene oar oh igre 2
2. Hind femur without a tubercle (Fig. 13) ... 7
— Hind femur with a ventral subbasal tubercle
covered with black setulae (Figs. 14-15). .. 3
3. Abdomen with yellow maculae on at least
2nd tergum; thorax entirely yellow pilose; ab-
dominal sterna without modified hairs ..... 5
— Abdomen with gray pollinose maculae on all
terga (Figs.1—2); mesoanepisternum and post-
alar callus black pilose. Hind femur entirely
DICK YAS Rett! ba aust een rae -|
4. Eye pilose; abdominal sterna without tufts of
specialized pile (Fig. 12); front yellow pilose
ee eg te Seer ee ee ne LEER ee Te A sibiricus
— Eye bare; 2nd and 3rd abdominal sterna with
apical submedial tufts of modified pile (Fig.
I) stront black pilose. =- -rae ee oe
5. Hind femur black on basal % or more; hind
femoral process elongate, with long setae
BR MS LO Se a I: A Sar fruterorum
— Hind femur yellow on basal % or more; hind
femoral process short, with short setae .... 6
6. Vertex broad, about as long as broad, black
pilose; hind trochanter with black setulae;
front tarsus with apical 2 tarsomeres dark;
metasternum bare divisus
— Vertex narrow, almost twice as long as broad,
yellow pilose on upper % or more; hind tro-
chanter yellow pilose, without black setulae;
front tarsus entirely orange; metasternum pi-
LOSE: AAA Eee ale flavifacies
7. 6th and most of 7th abdominal segments
shiny; vertex completely yellow pilose ... 14
— Genitalia segments grayish yellow pollinose;
VeLtexapantiallyablacks piloses-ssrielae eee ee 8
8. 2nd and 3rd terga without yellow maculae
(see couplet 11 for other characters) ... brooksi
porcus
636
2nd and 3rd terga with large yellow lateral
maculae
. Vertex narrow, at least twice as long as wide
posteriorly; ocellar triangle narrow; medial
ocellar distance much greater than distance
between lateral ocelli flavifacies
Vertex broader, about as long as wide poste-
riorly; ocellar triangle broader, medial ocellar
distance equal to or less than distance be-
tweenslaterdl:ocelliiees asain enn 10
. Front tibia extensively dark apically; hind fe-
mun black basallyaeeee tae at anaes 13
Front and middle tibiae yellow; hind basotar-
Somenre withoutiblackssctacyeee nse 11
. Hind femur black on basal %4, rarely very nar-
rowly yellow on base, with long pile baso-
posteriorly versicolor
Hind femur broadly yellow basally, yellow on
at least basal 4 or more, without any long pile
. Front yellow pilose; cercus elongate posteri-
if eel le Ere Bee Oe ee eine sare omen e laetus
orly
Front black pilose; cercus oval
. Mid tibia extensively dark apically; hind tibia
black on basal and apical %, yellow medially;
hind basotarsomere with a few black bristles
apicolaterally, without long lateral pile; front
partially or entirely yellow pilose
Mid tibia entirely yellow; hind tibia yellow
on basal *%, black apically; hind basotarso-
mere without black bristles, with long pile lat-
erally; front black pilose
rex
consimilis
. Mesonotum without distinct pollinose vittae;
hind femur narrow and arcuate; genitalia
large, with 6th segment as long as 4th tergum;
hind basotarsomere shorter pilose, with pile
only about as long as tarsal width
Mesonotum with distinct pollinose vittae;
hind femur broader and not arcuate; genitalia
smaller, with 6th segment only about 4% as
long as 4th tergum; hind basotarsomere lon-
ger pilose, with pile distinctly longer than tar-
sal width integer
. Hind femur entirely black; abdomen black,
with gray pollinose maculae on all terga;
mesonotum and mesoanepisternum extensive-
ly black pilose; fore and mid femora exten-
sively black pilose; hind basotarsomere with
black apicolateral bristles along anterior edge;
ocellar triangle broad, with medial ocellar dis-
tance much less than distance between lateral
ocelli porcus
Femora much more extensively yellow; ab-
domen partially yellow, at least with lateral
margins partially yellow; mesonotum and me-
soanepisternum yellow pilose ..........
. Abdomen with lateral margins broadly yellow
7h
20.
PV
2D,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and without or with very small medial yellow
maculae; femora extensively yellow, with
only a narrow dorsal black vitta on medial %;
ocellar triangle equilateral, with lateral ocel-
lus broadly separated from eye margin, sep-
arated by about twice its diameter, with me-
dial ocellar distance subequal to distance be-
tween lateral ocelli; hind tarsus without black
bristles: «> s:sces Sy. yee se cre eee brooksi
Abdomen with large yellow maculae at least
on 2nd and 3rd terga 117)
Hind femur yellow on basal % or more; ocel-
lar triangle larger, with lateral ocellus narrow-
ly separated from eye margin, separated by
about its) diameter r- sien eee ae a
Hind femur black or dark brown on basal %
or more, although base maybe narrowly yel-
low
. Fore tibia black apically, intense at least dor-
sally
Fore tibia entirely yellow, rarely slightly dark-
ened antero-ventrally
. Occiput dorsally with row of long black pile;
face profile straight; costa with golden pile
restricted basally, not extending to humeral
crossvein frutetorum
Occiput entirely yellow pilose; face profile
convex; costa with extensive golden pile ba-
sally, extending beyond humeral crossvein .
Fae nee eens Sedaris stele eee LS Ab versicolor
Mid tibia entirely yellow; hind tibia yellow
on basal % or more; fore basotarsomere with-
out black spinules; mesonotum without me-
dial vitta consimilis
Mid tibia black on apical % or more; hind
tibia black basally and apically, narrowly yel-
low medially; fore basotarsomere with black
spinules posteroventrally; mesonotum with a
narrow medial pollinose vitta
Front narrow, about ¥, of head width at ocellar
triangle; ocellar triangle approximately equi-
lateral, with medial ocellar distance about
equal to or greater than distance between lat-
eral ocelli; hind basotarsomere with a row of
short black bristles along anterior edge; hind
femur frequently black on apical % . . flavifacies
Front broad, about % or more of head width
at ocellar triangle; ocellar triangle broader,
with medial ocellar distance much less than
distance between lateral ocelli
rex
Ma Ee 22,
Hind basotarsomere with a double row of
short black bristles along anterior edge ....
Seisgslecd 3, Sua, Hite, hed Seas ake ne ore ge ee divisus
Hind basotarsomere usually without any
black bristles, at most with 2—3 black bristles
3. Hind basotarsomere with long erect pile, at
VOLUME 99, NUMBER 4
least laterally, with pile about as long as or
longenithanttansallwidth) 32 3-1 acne. -
— Hind basotarsomere with shorter appressed
integer
ILS Bei teres, hee ene chem iat lets wa teas) date toe 24
24. Mesonotum with pollinose vittae either absent
OFLODSCUTE MAES eS SAE eal eles obsoletus
Mesonotum with pollinose vittae distinct... 25
25. Hind femur entirely yellow anteriorly; fore
and mid tibiae entirely yellow
— Hind femur black medially; fore and mid tib-
lacwusuallyidarksbasalliys.-)- ae 4 ne
kurentzovi
laetus
CHECKLIST OF PARHELOPHILUS SPECIES
brooksi Curran. Alberta to Wisconsin.
Parhelophilus brooksi Curran 1927: 90 3
Manitoba, Petchipegosis (HT ¢ CNC).
Lunomyia pollinaria Fluke 1939: 373
62 Wisconsin, Spooner (HT 6
AMNH). Syn. Fluke 1953: 208.
consimilis Malm. Norway to western Si-
beria, south to France & Italy.
Helophilus_ consimilis Malm 1863: 80
Sweden, Stadsvassen. T ?
divisus (Loew). Michigan to Ontario, south
to Indiana & Florida.
Helophilus divisus Loew 1863: 316
(1864: 200 #4:78) District of Columbia
(ST 36 MCZ).
flavifacies (Bigot). Quebec, south to Mis-
sissippi & Florida.
Helophilus flavifacies Bigot 1883: 344 3
Maryland, Baltimore (HT ¢ BMNH).
Helophilus anniae Brimley 1923: 278 3
North Carolina, Raleigh (HT ¢
USNM). N. syn.
frutetorum Fabricius. Scandinavia to Sibe-
ria, south to Spain, Italy, Bulgaria & Kyr-
ghyzstan.
Syrphus frutetorum Fabricius 1775: 765
? England (T ? Unknown, not stated in
Fabricius)
Syrphus femoralis Fallén 1817: 31 ?
Sweden, “in paroecia Farhult’’ (T ZIL)
Helophilus frutetorum var. xanthopygus
Loew 1846: 149 Italy, Sicily, Syracuse
(T ? ZMHU)
integer (Loew). Ontario & Quebec, south
to North Carolina.
Helophilus integer Loew 1863: 314 ¢
New York (ST ¢ MCZ).
637
kurentzovi Violovitsh. Siberia, Korea, Ja-
pan.
Parhelophilus kurentzovi Violovitsh
1960: 207 3d 2 Russia, Sakhalin, Yuzh-
no-Sakhalinsk (HT @ ZISP).
Parhelophilus obscurior Violovitsh
1960: 209 3 2 Russia, Sakhalin, Yuzh-
no-Sakhalinsk (HT ¢ ZISP). N. syn.
Helophilus citricornis Shiraki 1968: 214
36% Japan, Oze. (HT 3 NIAS). N. syn.
laetus (Loew). British Columbia to Quebec,
south to New Mexico, Louisiana & North
Carolina.
Helophilus laetus Loew 1863: 315 3
New York & Wisconsin (ST ¢ MCZ).
Helophilus aureopilis Townsend 1895:
51 3 Michigan, Constantine (HT 6
UKaL). Syn. Hunter 1897: 139.
Parhelophilus currani Fluke 1953: 128
3 @ Louisiana, 15 miles east of Creole
(HT 3 UKaL). N. syn.
obsoletus (Loew). Alaska to Ontario and
Maine, south to British Columbia & Wis-
consin.
Helophilus obsoletus Loew 1863: 314 3
Hudsons Bay Territory (ST 6 MCZ).
porcus (Walker). British Columbia to New
Brunswick, south to Wisconsin & Mary-
land.
Eumerus porcus Walker 1849: 554 3 On-
tario, Hudson’s Bay, Albany River, St.
Martin’s Falls (LT ¢ BMNH here des-
ignated).
rex Curran & Fluke. British Columbia to
Ontario, south to Colorado & West Vir-
ginia.
Parhelophilus rex Curran & Fluke 1926:
234 32 Ontario, Macdiarmid & Al-
gonquin Park; New York, Lake
George, Northwest Bay (ST 6 CNC).
sibiricus Stackelberg. Russia (Karelia to
Eastern Siberia).
Pleskeola sibirica Stackelberg 1924: 25
d Russia, “‘Sibiria centralis, Obj-En-
isseljskij kanal. septen. et orientem
versus Enisseisk’”? (HT 3 ZISP). N.
comb.
versicolor Fabricius. Scandinavia to Sibe-
638
ria, south to Spain, Italy, Bulgaria & Kir-
ghiz.
Syrphus versicolor Fabricius 1794: 283
Germany (T ? Unknown, stated as
‘Dom. Smidt”’ in Fabricius)
Parhelophilus almasyi Szilady 1940: 65
Kazakhstan, Taldy-Kurgan, Panfilov
(T ? MNM (destroyed?))
NOTES ON THE KEY AND SPECIES OF
PARHELOPHILUS
Parhelophilus flavifacies is keyed in 2
couplets because the femoral tubercle can
be low and difficult to distinguish in some
specimens. The types of flavifacies and an-
niae were examined and found to represent
the same species.
Parhelophilus obscurior Violovitsh and
Helophilus citricornis Shiraki are syn-
onyms of kurentzovi. I have studied the ho-
lotype of citricornis and Barkalov (in litt.)
has studied the types of the Violovitsh spe-
cies.
Parhelophilus currani Fluke is only a
pale southern race of Jaetus, the few spec-
imens of which I have seen differ from
more northern ones in that the abdominal
maculae are expanded and united to form
fasciae.
Parhelophilus brooksi (Curran) is prob-
ably best considered a species of Lejops
(Lunomyia) as placed by Fluke on the as-
sumption that the lack of the apical calcar
on the hind tibia is a secondary loss. The
species is, however, included in the key as
this is the genus in which users will prob-
ably still identify brooksi.
Helophilus pilosus Hunter is a species of
Lejops (Lejops), new combination.
ACKNOWLEDGMENTS
I thank David Grimaldi, American Mu-
seum of Natural History, New York
(AMNH); Brian Pitkin, The Natural Histo-
ry Museum [formerly the British Museum
(Natural History)], London (BMNH), Ser-
gei Kuznetzov, Zoological Institute, Acad-
emy of Science, St. Petersburg (ZISP); N.
Fukuhara, Insect Taxonomy Laboratory,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
National Institute of Agricultural Sciences,
Tsukuba (NIAS); H. Schumann, Humboldt
University, Berlin (ZMHU); J. R. Vocker-
oth, Canadian National Collection, Ottawa
(CNC); Philip Perkins, Museum of Com-
parative Zoology, Cambridge (MCZ); and
Gunilla Stahls, Finish Museum of Natural
History, Helsinki (UZMH) for permission
to study material in their care. Other col-
lection acronyms used are: MNM for Mag-
yar Nemzeti Muzeum, Budapest; UKaL for
Snow Entomological Museum at University
of Kansas, Lawrence; USNM for National
Museum of Natural History (=United
States National Museum), Smithsonian In-
stitution, Washington, D.C.; and ZIL for
Zoological Institute, Lund. I also thank Drs.
Neal Evenhuis, Bishop Museum, Honolulu
(BBM); Manya B. Stoetzel, Allen Norr-
bom, and James Pakaluk, Systematic En-
tomology Laboratory, USDA, Washington,
D.C., and Beltsville, MD, for their critical
reviews of the manuscript.
I am very greatful to Frank D. Fee of
State College, Pennsylvania (FEE, personal
collection) and Chris T. Maier, Connecticut
Agricultural Experimental Station, New
Haven (CTM, personal collection) who not
only reviewed this manuscript but also pro-
vided many distribution records of porcus
from their personal field work as well as
detailed notes on the collection sites. Dr.
Harry D. Pratt, Atlanta, GA, also provided
details on the Vermont site.
LITERATURE CITED
Aldrich, J. M. 1905. A catalogue of North American
Diptera. Smithsonian Miscellaneous Collection
46(2), 680 pp.
Bigot, J. M. EF 1883. Diptéres nouveaux ou peu con-
nus. 22° partie, XXXII: Syrphidi (2° partie). Es-
peces nouvelles, no. I. Annales des la Societé En-
tomologique de France (6) 3: 61—88; 221-258;
315-356.
Brimley, C. S. 1923. Additional Syrphidae (Diptera)
from North Carolina, with descriptions of two
supposed new species. Entomological News 34:
277-279.
Cole, EF R. and Schlinger, E. I. 1969. The flies of
Western North America. xii + 693 pp. Berkeley
and Los Angeles.
VOLUME 99, NUMBER 4
Curran, C. H. 1927. Descriptions of nearctic Diptera.
Canadian Entomologist 59: 79-92.
Curran, C. H. and C. L. Fluke. 1926. Revision of the
Nearctic species of Helophilus and allied genera.
Transaction of the Wisconsin Academy of Sci-
ences, Arts and Letters 22: 207-281.
Fabricius, J. C. 1775. Systema entomologiae, sistens
insectorvm classes, ordines, genera, species,
adiectis synonymis, locis, descriptionibvs, obser-
vationibvs. [32] + 832 pp. Kortii, Flensburgi et
Lipsiae [=Flensburg and Leipzig]. [1775.04.17]
Fabricius, J. C. 1794. Entomologia systematica emen-
data et aucta. Secundum classes, ordines, genera,
species adjectis synonimis, locis, observationibus,
descriptionibus. Vol. 4, [6] + 472 + [5] pp. C. G.
Proft, Hafniae [=Copenhagen].
Fallén, C. E 1817. Syrphici Sveciae [Part]. Pp. 31—
42. Lundae (=Lund). [1817.05.20]
Fluke, C. L. 1922. Syrphidae of Wisconsin. Transac-
tion of the Wisconsin Academy of Sciences, Arts
and Letters 20: 215-253, pls. 5-6.
Fluke, C. L. 1939. New Syrphidae (Diptera) from
Central and North America. Annals of the Ento-
mological Society of America 32: 365-375, 1 pl.
Fluke, C. L. 1953. Some syrphid fly synonomy. En-
tomological News 64: 208-209.
Girschner, E. 1897. tiber die Postalar-Membran
(Schiippchen, Squamulae) der Dipteren. Illustrier-
te Wochenschrift fiir Entomologie 2: 534-539,
553-559, 567-571, 586-589, 603-607, 641-645,
666-670, 6 pls.
Goffe, E. R. 1944. Some changes in generic nomen-
clature in Syrphidae (Diptera). Entomological
Monthly Magazine 80: 128-132. [1944.07.04]
Goffe, E. R. 1945. Note on the type-species of some
genera of Syrphidae (Diptera). Journal of the So-
ciety for British Entomology 2: 276-279.
[1945.11.30]
Goot, V. S. van der 1981. De zweefvliegen van
Noordwest-Europa en Europees Rusland, in het
bijzonder van de Beneluz. Bibliotheck van der
Koninklijke Nederlandse Natuurhistorische Ver-
eniging 32, 275 pp.
Graenicher, S. 1910. A preliminary list of the flies of
Wisconsin belonging to the families Bombyliidae,
Syrphidae and Conopidae. Bulletin of the Wiscon-
sin Natural History Society 8: 32-44.
[1910.05.??]
Hackman, W. 1980. A check list of the Finnish Dip-
tera. Il. Cyclorrhapha. Notulae Entomologicae 60:
117-162.
Hull, EF M. 1949. The morphology and inter-relation-
ship of the genera of syrphid flies, recent and fos-
sil. Transaction of the Zoological Society of Lon-
don 26: 257—408, 25 figs. [1949.05.??]
Hunter, W. D. 1897. Contribution to the knowledge
of North American Syrphidae.—II. Canadian En-
tomologist 29: 121-144, pl. 5.
639
Johnson, C. W. 1914. Notes on inadequate locality
labels (Dipt.). Entomological News 25: 123-126.
Johnson, C. W. 1923. Collecting Diptera in Maine.
Maine Naturalist 3: 7-10. [1923.08.15]
Johnson, C. W. 1925. Fauna of New England. 15. List
of the Diptera or two-winged flies. Occassional
Papers of the Boston Society of Natural History
7(15), 326 pp., 1 fig.
Johnson, C. W. 1985. Bogs of the Northeast. xvi +
269 pp. University Press of New England, Hano-
ver & London
Kertész, K. 1910. Catalogus dipterorum hucusque
descriptorum. Vol. 7, 470 pp. Lipsiae, Budapestini
(=Leipzig, Budapest).
Loew, H. 1846. Helophilus. Stettiner Entomologische
Zeitung 7: 116-127, 141-150, 164-169.
Loew, H. 1863. Diptera Americae septentrionalis in-
digena. Centuria tertia. Berliner Entomologische
Zeitschrift 7: 1-55.
Malm, A. W. 1863. Anteckningar 6fver Syrphici i
Skandinavien och Finland, med sarskildt afseende
pa de arter och former, hvilka blifvit funna i G6-
teborgs och Bohus lan. Goteborgs K. Veternsk.-o.
VitterhSamh. Handl. 8: 1-81.
Miliczky, E. R. and E. A. Osgood. 1979. The effects
of spraying with Sevin-4-oil on insect pollinators
and pollination in a spruce-fir forest. Technical
Bulletin, Life Sciences Agricultural Experimental
Station of University of Maine 90, 21 pp.
Osburn, R. C. 1907. The Syrphidae of British Colum-
bia. Bulletin of the British Columbia Entomolog-
ical Society 8: 1—4.
Osburn, R. C. 1908. British Columbia Syrphidae, new
species and additions to the list. Canadian Ento-
mologist 40: 1-14, 1 fig.
Osten Sacken, C. R. von 1858. Catalogue of the de-
scribed Diptera of North America. Smithsonian
Miscellaneous Collections 3(1), vii-xx, 92 pp.
Osten Sacken, C. R. von 1875. A list of the North
American Syrphidae. Bulletin of the Buffalo So-
ciety of Natural Science 3: 38-71. [1875.11.??]
Osten Sacken, C. R. von 1878. Catalogue of the de-
scribed Diptera of North America. Smithsonian
Miscellaneous Collection, 270, xlvii + 276 pp.
[1878.10.??]
Peck, L. V. 1988. Family Syrphidae. pp. 11—230. In
Soos, A., ed., Catalogue of Palaearctic Diptera.
Vol. 8, Syrphidae—Conopidae. 363 pp. Akade-
miai Kiado, Budapest
Petch, C. E. and J. B. Malthais. 1932. A preliminary
list of the insects of the Province of Quebec. Part
II, Diptera, by A. E Winn and G. Beaulieu. Re-
vised and supplemented. Annual Report, Quebec
Society for the Protection of Plants 23/24 (supple-
ment 24), 100 pp.
Sack, P 1931. 31. Syrphidae [part]. Pp. 241—288. Jn
Lindner, E., ed., Die Fliegen der paladarktischen
Region. 4(6). Stuttgart [1931.04.29]
640
Shiraki, T. 1968. Syrphidae (Insecta: Diptera). Vol. 3,
[viii]+272 pp., 47 pls. In Fauna Japonica. Bio-
geographical Society of Japan, Tokyo.
[1968.03.30]
Stackelberg, A. A. 1924. Syrphidarium novorum pa-
laearcticorum diagnoses. Wiener Entomologische
Zeitung 41: 25-29.
Stackelberg, A. A. 1970. [49. Fam. Syrphidae—Hov-
er flies.] Pp. 11-96. In Bei-Bienko, G. Ya (ed.),
[Keys to the insects of the european part of U. S.
S. R.] 5(2) Leningrad [In Russian]
Stackelberg, A. A. 1988. 49. Fam. Syrphidae. Pp. 10—
148. In Bei-Bienko, G. Ya (ed.), Keys to the in-
sects of the european part of U. S. S. R. 5(2)
Smithsonian Institution Libraries & National Sci-
ence Foundation, Washington
Strickland, E. H. 1938. An annotated list of the Dip-
tera (flies) of Alberta. Canadian Journal of Re-
search (D, Zoological Sciences) 16: 175-219.
Szilady, Z. 1940. iiber palaarktische Syrphiden. IV.
Annales historico-naturales Musei nationalis Hun-
garici (Zoology) 33: 54—70.
Thompson, F C. 1997. The Biosystematic Database
of World Diptera. Diptera Data Dissemination
Disk 1. Also at URL: http://www2.sel.barc.usda.
gov/Diptera/biosys.htm
Thompson, FE C. 1997. A key to the genera of the
flower flies of the Neotropical Region with the
description of two new genera and eight new spe-
cies. Contribution of the American Entomological
Institute, 56 pp. [In Press]
Townsend, C. H. T. 1895. Contributions to the dip-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
terology of North America --I. Syrphidae. Trans-
action of the American Entomological Society 22:
33-55.
Verrall, G. H. 1901. Platypezidae, Pipunculidae, and
Syrphidae of Great Britain. Vol. 8, (vi) + 691 pp.,
457 figs. In his British Flies. London, Cambridge.
Violovitsh, N. A. 1960. New palaearctic Syrphidae
(Diptera) from the Far Eastern Territory of the
USSR. Entomologicheskoe Obozrenie 36: 748-—
755. [In Russian]
Violovitsh, N. A. 1983. Sirfidy Sibiri (Diptera, Syr-
phidae). 241 pp. Nauka, Novisibirsk. [In Russian,
see below for translation]
Violovitsh, N. A. 1986. Siberian Syrphidae (Diptera)
(Sirfidy Sibiri) Novosibirsk, 1983. Verslagen en
Technische Gegevens, Instituut voor Taxonomis-
che Zoologie (Zoologisch Museum), Universiteit
van Amsterdam. 288 pp. [Translation of the fore-
word and keys of Violovitsh 1983]
Walker, E 1849. List of the specimens of dipterous
insects in the collection of the British Museum.
Vol. 3, Pp. 485-687. London [1849.06.30]
Williston, S. W. 1887. Synopsis of the North Ameri-
can Syrphidae. Bulletin of the United States Na-
tional Museum 31, xxx + 335, 12 pls. (1886).
[Before 1887.06.30]
Wirth, W. W., Y. S. Sedman and H. V. Weems, Jr.
1965. Family Syrphidae. pp. 557—625. Jn Stone,
A., C. W. Sabrosky, W. W. Wirth, R. H. Foote,
and J. R. Coulson, eds., A catalog of the Diptera
of America north of Mexico. U. S. Department of
Agriculture, Handbook 276, 1696 pp.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 641-644
RUDOLFINA CAVERNICOLA, A NEW SPECIES OF CAVE-ASSOCIATED
SPHAEROCERIDAE (DIPTERA) FROM COLORADO AND ARIZONA
S. A. MARSHALL AND S. FITZGERALD
(S.A.M) Department of Environmental Biology, University of Guelph, Guelph, Ontario,
Canada NIG 2W1; (S.F) Department of Entomology, Colorado State University, Fort
Collins, CO 80523, U.S.A.
Abstract.—Rudolfina cavernicola, new species, is described from several collections
from Kremmer’s Cave, Colorado, and a single specimen from Arizona. Characters are
given to separate the two western North American Rudolfina. Six species of Sphaerocer-
idae are listed from Kremmer’s Cave, and the cave habitat is described.
Key Words:
The genus Rudolfina Rohaéek can be rec-
ognized on the basis of the stout, upturned
female cerci that are fused with the epiproct
and middle part of tergite 8; the long co-
stagial bristle; and the characteristic dark
lobes of male sternite 5. There is one de-
scribed Nearctic species, Rudolfina digitata
Marshall, and one described Palearctic spe-
cies, R. rozkosnyi (Rohacek), both of which
exhibit disjunct arctic-alpine distributions
(Marshall 1991). Rudolfina includes several
undescribed southeastern Nearctic and Neo-
tropical species, but Rudolfina cavernicola,
new species, is closely related to the west-
ern North American R. digitata and the Eu-
ropean R. rozkosnyi.
The cave-inhabiting Sphaeroceridae of
eastern North America were discussed by
Marshall and Peck (1984), who recorded 10
species of which at least one (Spelobia te-
nebrarum (Aldrich)), is a true troglobite,
and at least one, (Terrilimosina racovitzai
(Bezzi)), is a troglophile. Little is known
about cave Sphaeroceridae in western North
America, although one troglophilic species,
Limomyza cavernicola Marshall, was _ re-
cently described from in or near caves in
Missouri, Kentucky, Oklahoma, and Colo-
rado (Marshall 1997). Most of the type se-
Sphaeroceridae, Rudolfina, caves
ries of the new species described here, Ru-
dolfina cavernicola, is from Kremmer’s
Cave, Colorado. Spelobia pseudosetaria
(Duda), Spelobia tuberculosa Marshall,
Spelobia maculipennis (Spuler), Xe-
nolimosina sicula Marshall and Limomyza
cavernicola Marshall were collected in
Kremmer’s Cave with Rudolfina caverni-
cola.
KEY TO WESTERN NORTH AMERICAN
RUDOLFINA
— Eye height ca. 1.8 genal height. Second cos-
tal sector longer than third. Male sternite 5
with 3 posteromedial lobes, outer 2 small,
equal to one another (Fig. 4). Distal lobe of
surstylus setose, rounded (Fig. 1). Gonostylus
expandedspreapicallya (i lens) saa iene ee
ites, See ee Rudolfina cavernicola, new species
— Eye height ca. 1.4 genal height. Second cos-
tal sector shorter than third. Male sternite 5
with two posteromedial lobes, one bifid and
one single. Distal lobe of surstylus elongate,
bare, with multiple translucent lobes. Gono-
stylus narrow, tapered apically. ..........
Sas ie ciety Sue Se hee Glee Rudolfina digitata Marshall
Rudolfina cavernicola Marshall and
Fitzgerald, NEw SPECIES
(Figs. 1-6)
Description.— Body uniformly pale
brown; length ca. 2.0mm. Interfrontal plate
642 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-6. Rudolfina cavernicola. 1-4, Male abdominal structures. 1, Terminalia, left lateral. 2, Terminalia,
posterior. 3, Aedeagus and associated structures. 4, Sternites 5-7. 5—6, Female abdominal structures. 5, Tergites
7-9 and cerci. 6, Spermathecae.
equal in depth and width, bordered by 3 anteroventrally with a row of short, stout
interfrontal bristles, the lower pair cruciate bristles on distal two thirds; mid tibia of
over lunule. Eye height 1.8 times genal female with a single small anteroventral
height. Mid tibia of male slightly curved, bristle near middle. Dorsocentral bristles in
VOLUME 99, NUMBER 4
a single large prescutellar pair; 2 pairs of
prescutellar acrostichal bristles, middle pair
very long. Wing with costagial bristle long,
longer than dorsocentral bristle; second cos-
tal sector slightly longer than third, costa
ending 2—3 vein widths beyond apex of
Ryis.
Male abdomen: Sternite 5 with 3
posteromedial lobes (Fig. 4). Surstylus with
a broad, subquadrate basal part and a nar-
row, setose distal part (Fig. 1). Epandrium
with 3 long posterodorsal bristles on each
side (Fig. 2). Subanal plate large, broadly
fused; cerci small, setose. Gonostylus api-
cally expanded, boot-shaped (Fig. 3). Dis-
tiphallus broad, complex, functional ventral
surface with diverging arms, distal part of
functional dorsal surface with crenulate
plate on each side.
Female abdomen: Tergite 9 (Fig. 5) pro-
longed into a short, blunt lobe between the
2 halves of tergite 8, lobe dark and sepa-
rated from rest of tergite 9 by pale area,
main part of tergite 9 with a longitudinal
pale strip. Sternite 7 large, dark, posterior
margin spinulose; sternite 8 reduced, with
4 small, stout bristles. Sternite 9 postero-
medially thin and pale. Cercus distinctly
upturned, with a stout spur at apex (Fig. 5).
Spermathecae round, tire-shaped; stem
long.
Holotype.—¢d, COLORADO. Larimer
€o., 3 April, 1992, S. Fitzgerald, Krem-
mer’s Cave, 6000’, Rt 287,6 mi N of Jct.
Rt. 14. (Canadian National Collection)
Paratypes.— Colorado. Kremmer’s Cave,
Bros22 vit 1996 (16 ,2°9, 30:i1x—27-x.1996;
93,32 in pit trap @85’ from entrance; 12
in pit trap @ 30’ from entrance; none taken
in pit traps @ 9’ from entrance during same
period), Scott Fitzgerald (University of
Guelph and Colorado State University col-
lections). Arizona. Hospital Flat, Pinaleno
Mts., 8950’, 3.viii.1965, H. Leech (16, Cal-
ifornia Academy of Sciences).
Comments.—Although the male genita-
lia of R. cavernicola are strikingly different
from those of R. digitata, the female ter-
minalia of these closely related species are
643
very similar in structure. Both species have
a divided tergite 8, with the anterior part of
tergite 9 separate from the main part of ter-
gite 9 and between the halves of tergite 8.
However, tergite 8 of R. cavernicola is not
strongly wrinkled like that of R. digitata,
and the spermathecae are longer. The biol-
ogy of R. digitata is unknown, but since it
has smaller eyes than those of R. caverni-
cola it is assumed that both of these species
are subterranean in habit.
Kremmer’s Cave is a small solution cave
formed in Ingleside limestone along the
east slope of the hogback in northern Col-
orado (Parris 1973). The cave is basically
one 100 ft. long room sloping downward at
about 35°. The room is largest medially,
about 20 ft. wide with a 7 ft. ceiling, and
narrowest near the entrance and low, ter-
minal crawlway. The pit entrance is about
2 ft. in diameter and 3 ft. deep, and al-
though the cave is short, it becomes cave-
like (dark, humid, and of a relatively con-
stant temperature) quickly, due to the small
opening to the outside. In August the am-
bient air temperature taken at 25 ft. in from
the bottom of the pit entrance was 42° F
Specimens of R. cavernicola were col-
lected within the first 25 ft. from the bottom
of the pit entrance, henceforth referred to as
the entrance-way, using an aspirator, and at
30 and 85 ft. from the bottom of the pit
entrance with pitfall traps baited with slices
of ripe banana. Since R. cavernicola was
collected in sympatry with several other
species of sphaerocerids, field identification
was not possible and thus, the following
general observations include all above men-
tioned species. In the entrance-way flies
were observed resting or running on pieces
of moist wood, among broken rock, rock
pillars (probably reminiscent of the cave’s
discovery during a limestone mining oper-
ation (Parris 1973)), and on cave walls.
Flies jumped or ran when disturbed, but
were never observed to fly.
Rodent droppings, most covered with
various microfungi, seem to be the most
abundant food source in Kremmer’s cave.
644
However, the nature of the pit entrance al-
lows rain water to wash in organic debris,
a minimal amount of bat guano exists near
the back of the cave, and a rodent jaw bone
suggests at least an occasional carcass, all
of which may be possible larval food
sources. As most cave-associated sphaero-
cerids utilize a wide range of breeding sub-
strates (Papp and Plachter 1976), and R.
cavernicola was apparently attracted to ba-
nana, it is probable that R. cavernicola is
not restricted to a single food source.
ACKNOWLEDGMENTS
Thanks to Rebecca Langstaff for prepar-
ing the illustrations, and to Alison Foley for
help with cave measurements and the col-
lection of traps.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
LITERATURE CITED
Marshall, S. A.
primitive Limosininae (Diptera: Sphaeroceridae),
with five new species from United States, Mexico,
and Central America. Proceedings of the Ento-
mological Society of Washington 99:279-289.
Marshall, S. A. 1991. Rudolfina digitata sp.nov., a
new nearctic sphaerocerid with a disjunct alpine-
arctic distribution. Canadian Entomologist 123:
621-626.
Marshall, S.A. and S.B. Peck. 1984. Distribution of
cave-dwelling Sphaeroceridae (Diptera) of North
America. Proceedings of the Entomological So-
ciety of Ontario: 115: 37-41.
Parris, Lloyd E. 1973. Caves of Colorado. Boulder,
Colorado, Pruett Publishing Co., 247 pp.
Papp, L. and Plachter, H. 1976. On cave dwelling
Sphaeroceridae from Hungary and Germany. An-
nales Historico Naturales Musei Nationalis Hun-
garica 68: 195-207.
1997. Limomyza, a new genus of
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 645—660
A REVIEW OF THE GENUS HOPLISOIDES GRIBODO
(HYMENOPTERA: SPHECIDAE: GORYTIND IN NORTH AMERICA
RICHARD M. BOHART
Department of Entomology, University of California, Davis, CA 95616, U.S.A.
Abstract.—The 30 species of Hoplisoides known from the continent of North America
and its associated islands are described and keyed. Pertinent illustrations are given. Four
new species are presented: H. elotae (Sinaloa, Mexico), H. niger (Puerto Rico), H. sub-
costalis (Panama), and H. parkeri (Mexico). New synonyms are: knabi Rohwer 1911
provisionally = costalis Cresson 1872; maculipennis Cameron 1890 = iridipennis F. Smith
1856; umbonicida Pate 1941 = vespoides FE Smith 1873. New status are: hypenetes
Handlirsch 1895 as synonym of denticulatus Packard 1867; birkmanni Baker 1907 as
synonym of nebulosus Packard 1867; pygidialis W. Fox 1896 as full species, not subspe-
cies of costalis Cresson 1872.
Key Words:
Hoplisoides Gribodo is one of the largest
genera in the tribe Gorytini with about 70
species in the World. The species of North
America were keyed by W. J. Fox (1896),
who recognized 9, but a little over 100
years later 30 are now known. Among the
rather numerous characters of the tribe
Gorytini, the following combination will
identify Hoplisoides: forewing pictured,
hindwing media diverging close to cross-
vein cu-a, omaulus and acetabula present,
sternaulus at least partly present, propo-
deum punctate but without a spiracular
(stigmatal) sulcus, propodeal enclosure usu-
ally with at least traces of longitudinal car-
inula, T-I not petiolate or pedunculate, male
sterna V—VI with concealed hairbrushes.
Material studied has been about 1,000
specimens, including types, in some 30 mu-
seums in America and Europe. Muséums
are listed in the Acknowledgments. This is
in addition to the more than 1,500 speci-
mens in the Bohart Museum, which has the
largest and most complete collection of
North American species.
Hoplisoides, Sphecidae, Gorytini, North America, key
Hoplisoides wasps are ground-nesting,
and provision their nests with Cicadellidae
or Membracidae. A detailed summary of bi-
ology was given by Bohart and Menke
(1976:520):
ACKNOWLEDGMENTS
The following museums and their man-
agers have been most helpful in allowing
examination of their holdings, and have
kindly sent types when requested. The mu-
seums are identified by the pertinent city in
capitals. Many museums not listed have
sent specimens for identification which
have helped fill out distributional data.
Academie de Sciences de
Cuba). (i024. spomaseatee HAVANA(I)
Academy of Natural
SCIENCES esa EBs : PHILADELPHIA
American Museum of
Natural History ........ NEW YORK
California Academy of
SGlENnCeSt.sh .40neees SAN FRANCISCO
Canadian National
Collectionvay- Besse eek. OTTAWA
646
GoriellsUiniversityer, os. son ITHACA
Entomologisches Institut,
Technische Hochschule ...... ZURICH
Gundlach Collection,
Guba: iat. hae. oe. acres HAVANA(II)
Humboldt Museum: a.-%42 -o. BERLIN
Laval University Provancher
Collection: 234355 eS ee QUEBEC
Los Angeles County
Museum) 2-5 ee LOS ANGELES
Museum of Comparative
Zoology, Harvard CAMBRIDGE
Museum d’ Histoire Naturelle,
Switzerland. “so ee ek GENEVA
National Museum of
Natural History ..... WASHINGTON
Natural History Museum,
UR ng ee rete A a Oe LONDON
Naturhistorisches Museum of
PuStrl dy 24 coals see ee ee VIENNA
Utah* State University. 2. --.2..- LOGAN
Universitets Zoologiske
Museum 252026 24.5-) COPENHAGEN
University of California
Bohart Museuin.......2 2.026 05 < DAVIS
University of California
Essie Museums (s: : ir s4.. BERKELEY
University of Halle, Germany HALLE
University of Lund, Sweden ..... LUND
University of Nebraska State
VENER Sorted tae LINCOLN
University of California
Riverside Museum ...... RIVERSIDE
In the following key an important char-
acter is the nature of the metapleuron, its
shape, size of its lower pit (called antero-
ventral metapleural pit by Bohart and Men-
ke 1976:fig 3), and its punctation (or lack
thereof). The abbreviations MOD (median
ocellus diameter), F (flagellomere), T (ter-
gum), S (sternum), and PD (puncture di-
ameter) have been used in the keys and de-
scriptions below. An interesting condition
of most Hoplisoides males, and the result
of considerable synonymy, is the presence
of only 6 visible terga, instead of usual 7
in most gorytins, followed by an exserted
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
S-VIII as a pseudosting. Females are easily
distinguished by having a pygidial plate on
T-VI, and 12 antennal segments instead of
13 in the male. In general the terminology
of characters follows that described in Bo-
hart and Menke 1976.
KEY TO THE SPECIES OF HOPLISOIDES IN
NORTH AMERICA
Males
1. Mesopleuron impunctate or nearly so (some-
times with a few very fine punctures) ..... i
— Mesoplenronipunctates... -)2j-t ieee 6
2. ESleunbandedh 1%. 2 ahh. 2\ ae eee 3
— T-III banded with white or yellow ....... a
3. Markings yellow, propodeal enclosure with
carinulae incomplete posteriorly; Cuba
ater (Gmelin)
— Markings white, propodeal enclosure with
complete carinulae; Hispaniola . . . . alaya (Pate)
4. Propodeal enclosure carinulate, scutum and
propodeum (posteriorly) plainly punctate .. .
Sean Suye cit adsgece Chew eee iridipennis (F. Smith)
— Propodeal enclosure essentially smooth, pro-
podeal punctation various
5. Scutum well punctured, propodeum black and
with obvious punctures posteriorly, subdis-
coidal and second discoidal cells clear (based
on female characters); se. U.S. and Argentina
SES gterbis 3 Ghd semipunctatus (Taschenberg)
— Scutum and posteriorly red propodeum with
microscopically fine punctures, subdiscoidal
and second discoidal cells stained brownish
glabratus R. Bohart
6. S-III to V and venter of mesothorax with
dense, white, woolly pubescence, sometimes
bloomlike, S-V often with a tooth or carina
visible laterally: 7.2 ste. = 1s eee ee 7
— S-III to V without such pubescence, no tooth
ONS Willer oia: Aa. Ta eee 12
7. S-V with at most a lateral denticle ....... 8
— S-V with a definitely raised carina laterally
ea ee ee loa a icle cian By 4 10
8. Metapleural lower pit about as large as mid-
ocellus, F-VI-VIII strongly nodose beneath
Sele AEE EMER Es Ph ey fe oe floridicus R. Bohart
— Metapleural lower pit smaller than midocel-
lus; P-VI-VIE various, ... co.. «see 9
9. F-VI to VII strongly thickened and nodose
beneath, metapleuron usually at least partly
red, pubescence of sterna bloomlike
chee 2 eRe « seed seek cazieri R. Bohart
— F-VI-VII a little thickened beneath, but not
nodose; metapleuron all black; pubescence of
sterna 1.0 MOD or more long laterally ....
ois Rie gt Reena eae Ae denticulatus (Packard)
VOLUME 99, NUMBER 4
10. Scutal punctures fine, mostly with diameters
USs
16.
18.
less than 0.25 MOD, hindfemur mostly yel-
low outwardly, metanotum practically im-
PUNCctate Hee es eae projectus R. Bohart
Scutal punctures moderate, many with diam-
eters equal to 0.5 MOD, hindfemur either ex-
tensively brown or red outwardly
. T-V carina rounded across top, hindfemur
mostly red outwardly ..... confertus (W. Fox)
T-V carina flat across top, hindfemur mostly
brown outwardly ........ carinatus R. Bohart
. Metapleural lower pit smaller than midocellus
RPS ALLIS tein Sis cad st Actin SS eae ceer eee: 13
Metapleural lower pit about as large as or
larcemthansmidocellus) 24) eee eee 20
. Mesopleuron dentate below, metapleuron at
middle narrower than 1.0 MOD, body exten-
Sivielyenedsandhy ellOw, ee) ees r-n skeen 14
Mesopleuron not dentate below, metapleuron
at middle broader than 1.0 MOD, body vari-
QUSP ay as a se eck) eel tee ere 15
. Propodeum in posterior view red and yellow
Pei ros ea a ee diversus (W. Fox)
Propodeum in posterior view all or nearly all
eG gfe ed ay tee Se eee dentatus (W. Fox)
Forewing with black band from wing base to
apex of marginal cell, propodeal area just be-
yond metapleuron with series of diagonal
ridges (based on female); Panama .......
PP eins =, Sree ts ck pene subcostalis R. Bohart
Forewing without a back band from base of
wing to apex of marginal cell, propodeal area
just beyond metapleuron irregularly sculp-
tured or punctate
F-I at least 2 as long as greatest breadth,
metapleuron punctate, subdiscoidal and sec-
ond discoidal cells each with an apical dis-
CLEle ClOUG poe Sx 4 cin): Seog eel ere 17
F-I 1.2 to 1.7X as long as greatest breadth,
metapleuron not punctate, above cells not
Clouded) 2... 5b ysqlleus seceatce sieps a ot ee 18
. T-ITE-IV all or mostly dark, frons with punc-
tation partly obscured by silvery pubescence
spendidulus (Bradley)
T-IN_IV broadly yellow-banded (as on II, V, &
VI), frons with dense punctation not obscured
by pubescence; Mexico....... elotae R. Bohart
Propodeum usually and legs extensively red,
T-VI-VIHUI normally visible, metapleuron ta-
pering to a point below ..... tricolor (Cresson)
Propodeum and legs not extensively red,
spinelike S-VII protruding from end of ab-
domen, metapleuron below middle mostly
broader than 1.0 MOD
. Last few terga broadly black, yellow bands
en ES Ea, Sees Fe costalis (Packard)
Last few terga broadly yellow
BP TAS RWIS lel Slat SIe pygidialis (W. Fox)
|
i
i)
tN
ee)
N |
647
T-I-II-IV yellow banded, T-III-V—VI black,
Cuba
T-III yellow banded
. Propodeal enclosure with carinulae quite
weak or absent posteriorly; Cuba ........
insularis (Cresson)
Propodeal enclosure with carinulae all well
developed=3Cubaty ass eer xerophilus Alayo
. Scutum and mesopleuron with off-silvery or
pale golden pubescence, T-II unusually broad,
clypeus black vespoides (FE Smith)
Scutum and mesopleuron without unusual pu-
bescence, clypeus not usually all black ... 23
. Front and middle tibiae entirely yellow, ped-
icel bright yellow in front, yellow band of S-II
mostly impunctate; Cuba ....... Jibacoa Alayo
Front and middle tibiae not all yellow, pedicel
not bright yellow dorsally, yellow band of
SoU plainlyapunctate sa. opr ie en ra 24
. Clypeus transversely bent toward apex (bev-
Sled) EAS see Shae ter ae, Maen nd Wkettes, edo. 25
Clypeus not transversely bent toward apex 26
. Clypeus sharply beveled all across, propodeal
area alongside enclosure closely punctate
A Ae ee Os ae es hamatus (Handlirsch)
Clypeus rounded toward middle, propodeal
area alongside enclosure somewhat polished,
ablew pPuUNnchiTesses ars punctifrons (Cameron)
. T-I-II both coarsely and rather closely punc-
tured, PD = 4%—-% MOD .... parkeri R. Bohart
T-I-II or at least T-I not closely and coarsely
punctured Dia MOD Re eer 27
. T-III to VI all or nearly all black, propodeum
(excepienclosune) mmosthyarcdi renee
KCL ROI SR CISE Ee placidus (F. Smith)
T-III-IV (at least) yellow banded, propodeum
NOt-ed! era. wsercacttycsteAcs ied eos Eee eee 28
. Submarginal cell I, discoidal cell I, and apex
of medial cell all lightly clouded
i tar owen Soe ea nae BET placidus nebulosus (Packard)
Submarginal cell I, discoidal cell I, and me-
dial cell practically clear (Mexican specimens
usually with black lower edge on clypeus)
Da sg ok OW ct spilopterus (Handlirsch)
Females
. Mesopleuron impunctate or with only micro-
SCOPIC, PUNCHIRES Hs). seta cree Gy eres <b ereus. 2
Mesopleuron punctate) 2.42.2... q
> Teimbanded ee sok Se ISS Ce 3
T-III banded with white or yellow, at least
latenallyaRer yor ete tie Skene ote SES 5
. Thorax and abdomen almost all black, T-II
with traces of an apical yellow band; Puerto
RICO an. woken ers Sas se. niger R. Bohart
Thorax and abdomen not nearly all black, T-I
and T-IV with distinct bands
648
La:
16.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
. Markings yellow, propodeal enclosure not
carinulate posteriorly; Cuba ..... ater (Gmelin)
Markings white, propodeal enclosure com-
pletely carinulate; Hispaniola ..... alaya (Pate)
. Propodeal enclosure completely carinulate
iridipennis (Cameron)
Propodeal enclosure smooth
. Antenna all yellow, frons mostly yellow,
metanotum not yellow..... glabratus R. Bohart
Antenna black above, frons with yellow lat-
erally, metanotum yellow; se U.S., Argentina
Oat ad aes & eoit dae semipunctatus (Taschenberg)
. Metapleural lower pit smaller than midocellus
Metapleural lower pit about as large as or
larger than midocellus
. F-I 3X or 4X as long as greatest breadth .. 9
F-I 2X to 2.7 times as long as greatest breadth
. F-I 3X as long as greatest breadth, inner eye
margins nearly parallel below, no continuous
yellow or white band across summit of head,
subdiscoidal cell without a discrete apical
spot
F-I 4x as long as greatest breadth, inner eye
margins slanting slightly inward below, yel-
low band on summit of head continuous all
across, subdiscoidal cell with a discrete apical
spot
. Body mostly red, propodeum red in posterior
view, tergal yellow bands present on I-III (at
INOSE) vee. Meee 48. ea eee dentatus (W. Fox)
Body red and yellow, propodeum mostly yel-
low in posterior view, tergal yellow bands on
I Vae(atiledst) eee ee diversus (W. Fox)
Legs mostly brown; Mexico. . elotae R. Bohart
Legs mostly red spendidulus (Bradley)
. Metapleuron broader than 1.0 MOD above
but tapering to a point below, scutum and
mesopleuron mostly red ... . tricolor (Cresson)
Metapleuron broader than 1.0 MOD for most
of its length, scutum and mesopleuron various
1 che nike Se Ne eh Slee «SL epee rae 13}
. Scutum and mesopleuron mostly red, terga
mostly: yellow: «treet tes eu cee se 14
Scutum and mesopleuron mostly black, terga
VablOUS Fins aiacus cic Grace cions opty rease oboe 15
confertus (W. Fox)
Pygidium yellow or at least basally yellow
CAME Sea aer OOM PTAC ER eee one carinatus R. Bohart
. T-VI densely punctured, punctured part about
1.3X as long as broad at base ......... 16
T-VI punctured part not densely so, about
1.9—2.2X as long as broad at base ...... N7/
Legs mostly red, mesopleural punctures fine
Sey eee ig eee ae projectus R. Bohart
Legs mostly brown, mesopleural punctures
moderate’? Seta BARe awe ee. oe cazieri R. Bohart
. Forewing with black band along costal mar-
gin to end of marginal cell, hindwing with
dark cloud in basal two-thirds of medial cell;
Panamal (2 een eae subcostalis R. Bohart
Forewing without black costal band, hind-
wing not clouded
. T-III to V with narrow yellow bands, flagel-
lum partly reddish beneath, pygidial carinae
nearly parallel above, (T-I rarely all black)
Hh en OE GS Fae eee costalis (Cresson)
T-III to V broadly yellow, flagellum exten-
sively pale beneath, pygidial carinae gradu-
ally broadening above ..... pygidialis (W. Fox)
. T-III-IV nearly all dark, T-V—VI mostly yel-
low, propodeal enclosure with longitudinal
carinulae disappearing posteriorly; Cuba .. .
SS A Se ae PIS 3s ae Jibacoa (Alayo)
Terga not marked as above, propodeal enclo-
sure caninulactcomplete tse in er eee 20
20. Subdiscoidal cell with discrete black spot api-
Cally Wit hae ck ns ee ee 21
Subdiscoidal cell without discrete spot api-
Cally eh en oh ai Sen yc ae 24
. Propodeum black
. T-III all black, or nearly so, rarely with pale
marks on T-III-IV but metapleural lower pit
round! 2). 2a. Ae. See 22
T-III to T-V usually pale banded, metapleural
lower pit oval
. T-II black, propodeum mostly dark red; Cuba
SE reser monet AL Akt jJaumei (Alayo)
T-II pale banded, propodeum mostly orange
TOG 008 tis oh oy nt gins ee aes punctifrons (Cameron)
aE, Be hamatus (Handlirsch)
Propodeum (except enclosure) mostly red
oA IE Te Fata eee spilopterus (Handlirsch)
24. Pygidium with lateral carinae somewhat bent
25:
26.
Dike
28.
toward middle, T-II moderately punctate,
many punctures more than 1.5 PD apart or
more; Vil not mostly yellow ss ane 25
Pygidium with lateral carinae evenly curved,
othemcharacters!) vatlous# ee eee 26
T-I1I to VI all or practically all black, pro-
podeum (except enclosure) largely red
Re Be: Ge AT, ee placidus (F Smith)
T-IIl to T-V with apical yellow bands, pro-
podeum often all black
a EASA oe eee placidus nebulosus (Packard)
T-III black or nearly so (rarely partly banded
in xerophilus)
T-III banded
Propodeal enclosure weakly carinulate, be-
coming polished posteriorly, terga weakly
punctured; Cuba insularis (Cresson)
Propodeal enclosure completely carinulate,
terga well punctured; Cuba .. .xerophilus Alayo
Scutum and mesopleuron with off-silvery or
VOLUME 99, NUMBER 4
golden pubescence, sometimes bloomlike,
omaulus not continuous with sternaulus
a RES RO: ea eee vespoides (FE Smith)
— Scutum and mesopleuron without such pu-
bescence, omaulus smoothly continuous with
Stemauhuse § ajs 2600-2 Scream 29
29. T-I-II coarsely and rather closely punctured,
punctures averaging about 1.0 PD apart,
T-VL-_VII mostly or all yellow, submarginal
cell 3 clouded over forward one-third .....
> he & es iG) One eR ARE parkeri R. Bohart
— T-I not evenly and coarsely punctured, T-I
moderately so, other characters various ... 30
30. Mesopleural punctures adjacent to metapleu-
ron medium to fine, T-II black before apical
band, submarginal cell 3 clouded over about
Wneilifies Ss aes: «tba yaeeare pried oie denticulatus (Packard)
— Mesopleural punctures adjacent to metapleu-
ron coarse as on rest of mesopleuron, T-II of-
ten extensively orange red in front of apical
yellow band, submarginal cell 3 clouded over
at least two-thirds ........ floridicus R. Bohart
Hoplisoides alaya (Pate)
Psammaecius alaya Pate 1947:96. Holo-
type 6, “San Domingo” (PHILADEL-
PHIA).
Among the 6 species with a nearly im-
punctate mesopleuron, alaya is the only one
with a combination of white bands on I-II—
IV, and complete carinulae on the propodeal
enclosure. This small species has the me-
tapleural lower pit larger than the mid-
ocellus, and T-II is finely punctate. T-V is
distinctly punctured.
The 5 6 and 4 @ I have seen were all
collected on Hispaniola.
Hoplisoides ater (Gmelin)
Crabro tricinctus Fabricius 1775:375.
‘‘America’’. Lectotype d designated by
van der Vecht 1961:49 (COPENHA-
GEN).
Vespa ater Gmelin 1790. New name for
Crabro tricinctus Fabricius 1775:375,
preoccupied by Vespa tricincta Fabricius
1775:363 (now in Sphecius).
Vespa tristrigata Fabricius 1794:459. Lec-
totype 2, ““American Islands”’ designated
by van der Vecht 1961:49.
Lestiphorus behni Dahlbom 1842:11, Ho-
649
lotype 2 (LUND). Synonymy by Dahl-
bom 1845:483.
Harpactus scitulus Cresson 1865:147. Ho-
lotype 2, Cuba (HAVANA-ID. Synony-
my by Dalla Torre 1897:555.
Although related to alaya, the yellow
markings and posteriorly incomplete cari-
nae of the propodeal enclosure are differ-
entiating. I know the species only from 4
3 and 3 @ from Cuba. The female abdo-
men was figured by Alayo (1969). I have
seen the lectotype in the Copenhagen Col-
lection.
Hoplisoides carinatus R. Bohart
Hoplisoides carinatus R. Bohart 1968:287.
Holotype 6, Madera Canyon, Santa Cruz
Co., Arizona (DAVIS).
This species is one of six that have wool-
ly pubescence on S-III to VI of males.
Three of these have an obliquely placed ca-
rina laterally on S-V. These are carinatus,
confertus, and projectus. In confertus the
carina is relatively high (about 2.0 MOD),
in projectus lower (about 1.5 MOD) and in
carinatus lowest (about 1.0 MOD). All
three have the metapleural lower pit smaller
than a midocellus, and the male flagellum
only moderately swollen on F-VI-VII. In
carinatus the punctures of the frons below
the ocelli are larger, deeper, and closer than
those of the other two species. Characters
given for the females in the key hold fairly
well.
Distribution records are from southern
California (Temecula, Riverside, Big Pine),
southern Arizona (Phoenix, Tucson, Pearce,
Continental, Bowie, Sahuarita, Madera
Canyon, Portal, Nogales), New Mexico
(Rodeo); Sonora, Mexico (Cocorit, Mag-
dalena, Santa Ana), and Chihuahua, Mexico
(near Chihuahua).
Hoplisoides cazieri R. Bohart
Hoplisoides cazieri R. Bohart 1968:288.
Holotype 6, Carr Canyon, Huachuca
Mts., Cochise Co., Arizona (NEW
YORK).
650
The three Hoplisoides with woolly S-III
to VI in the male but no definite carina on
S-V are cazieri, denticulatus, and floridicus.
The male of cazieri differs from floridicus
by its much smaller metapleural pit, and by
its smaller pleural punctures overall. From
denticulatus it differs by its more nodose
form beneath F-VI-VII. Also, cazieri males
have no denticle laterally on S-V, whereas
denticulatus males usually have a percep-
tible one. Females of the three species are
not so easily separated, but characters given
in the key should suffice. An interesting
feature of both sexes is the partial or com-
plete redness of the metapleuron. This is of-
ten associated with a red streak diagonally
across the mesopleuron, and an extension
of the yellow on T-I.
Distribution records are from Arizona
(Huachuca and Chiricahua Mts.), Mexico
(Jalisco, Durango, Morelos, Oaxaca, Chia-
pas), Nicaragua (Chinandega), and Costa
Rica (Canfas). I have studied 41 ¢ and 4 @.
Hoplisoides confertus (W. Fox)
Gorytes confertus W. Fox 1896:525. Lec-
totype £ (seen) designated by Cresson
1928:47, ‘“‘Montana’’ (PHILADEL-
PHIA).
Gorytes imperialensis Bradley 1920:118.
Holotype d (seen), Brawley, California
(ITHACA). Synonymy by Bohart in Bo-
hart and Menke 1976:320.
The relatively high carina laterally on
S-V of the male sets this species apart from
its relatives, carinatus and projectus. In ad-
dition the length of the pubescence on S-VI
of the male (2 to 3 MOD) is remarkable.
The female is difficult to separate from that
of carinatus, but the partly or all yellow
pygidium of the latter is helpful. Females
of both species have the metapleural lower
pit smaller than is the case with floridicus.
Also, the extensive red markings of the tho-
rax differentiate confertus and carinatus
from denticulatus, projectus, and cazieri.
Distribution includes California (eastern
and southern), New Mexico (Tornero), Col-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
orado (Hasty, Kit Carson), Texas (Llano
Co., Alpine, Canyon, Santa Elena Canyon,
Randall Co.), Oklahoma (Buffalo), Kansas
(Clay Co., Scott Co., Lakin), ‘““Montana’’,
Nebraska (Chadron), Mexico (Samalayuca,
Chihuahua, Saltillo).
Hoplisoides costalis (Cresson)
Gorytes costalis Cresson 1872:225. Holo-
type 2 (seen), Texas (CAMBRIDGE).
?Gorytes knabi Rohwer 1911:569. Holo-
type od (seen), Progreso, Yucatan
(WASHINGTON).
Tentative new synonym.
Bohart and Menke (1976) listed pygidi-
alis as a subspecies of costalis but I now
consider it to be a valid species based on
distribution and markings. At the same time
they gave knabi species status. An exami-
nation of the holotype of knabi, kindly sent
by A. S. Menke, leads to the doubtful syn-
onymy above. In body structure, including
the tiny metapleural lower pit, relatively
simple flagellum, stout F-I, wing clouding,
black clypeus, and moderately coarse punc-
tation, knabi is similar to other males of
costalis | have seen. However, on the type
of knabi the yellow band on the metanotum
and large yellow area on the propodeum
posteriorly, are quite unusual.
Characteristics of costalis are a small
metapleural lower pit, a short male F-I
(about 1.3 as long as broad), a silvery pu-
bescent male clypeus which is often all
black, a coarsely punctured propodeum (ex-
cept enclosure), a moderately punctured
T-II, and a long narrow female pygidium
(about 2X as long as broad).
The distribution is east of the 100th me-
ridian, from New York to Nebraska and
south to Missouri and Florida. Mexican lo-
calities are Vera Cruz (Cordoba), Tamauli-
pas (Sierra Picachoa), and Hidalgo (Acto-
pan). H. knabi was from Yucatan (Progre-
SO).
Hoplisoides dentatus (W. Fox)
Gorytes dentatus W. Fox 1893:116. Lecto-
type d (seen), designated by Cresson
VOLUME 99, NUMBER 4
1928:47, Grand Canyon, Arizona (PHIL-
ADELPHIA).
The tooth on the lower mesopleuron is
the most distinctive feature of this species
as well as of diversus. Other characters
shared by the two species are an extremely
narrow metapleuron, an angled male omau-
lus, a short male F-I but female F-I 3X as
long as broad, T-VII usually visible in
males. Since both species occur in Califor-
nia, and Baja California Sur, Mexico, they
may be conspecific. However, there seems
to be a constant difference in markings, so
I have kept them separate. Both sexes of
dentatus have the propodeum red with
sometimes a faint yellowish suggestion.
Also, females have yellow tergal bands on
I-II or rarely I-III. Females of diversus
have the terga much more extensively yel-
low. Similarly, males have well formed yel-
low bands on I to III only.
I have seen 12 ¢ and 10 @ from Cali-
fornia (Antioch, near Pearblossom, Jacum-
ba), Arizona (near Eloy, near Sentinel),
New Mexico (Las Cruces), and Mexico:
Baja California Sur (near San Ignacio).
Hoplisoides denticulatus (Packard)
Gorytes denticulatus Packard 1867:430.
Holotype ¢ (not female) (seen), ‘‘Loui-
siana’”’ (PHILADELPHIA).
Gorytes barbatulus Handlirsch 1888:408.
Syntype 6, & (studied); d, Illinois, Tex-
as (GENEVA), 2, New Orleans, (ZU-
RICH). Synonymy by Bohart in Bohart
and Menke 1976:521.
Gorytes hypenetes Handlirsch 1895:894.
Syntype d (seen) “Columb” and ‘‘mex-
icanus Laguaira’’ (La Guaira, Venezue-
la?) (BERLIN). New status.
I have studied Packard’s type. His name
apparently referred to the uneven male fla-
gellum rather than to the denticle on S-V,
as might be supposed. The species is related
to cazieri and floridicus, both of which have
S-III to V in males with woolly pubescence
but no carina on S-V. Most males of den-
ticulatus have a discernible denticle later-
651
ally on S-V. Characteristics of the species
are the metapleural lower pit (slightly
smaller than a midocellus), black T-II basad
of yellow band, mesopleural punctures be-
coming 2—3 PD apart toward metapleuron.
Yellow bands on T-IV—V are usually thin
to moderate, but some 5 pair from Vera
Cruz, Mexico are extensively yellow on
T-IV to VI.
The distribution, based on 80 ¢ in the
Bohart Museum collection, is mostly east
of the 100th meridian (Florida, Missouri,
Georgia, Illinois, Texas, Oklahoma, Ne-
braska, Colorado). Most Mexican states are
represented as well as Costa Rica, El Sal-
vador, and Venezuela.
Hoplisoides diversus (W. Fox)
Gorytes diversus W. Fox 1896:527. Syntype
3, 2 (seen), Los Angeles, California
(WASHINGTON).
See discussion under dentatus and char-
acters given in male and female keys.
I have studied 10 ¢ and 4 2 from Cali-
fornia (Cawelo Jct. in Kern Co., Beaumont,
Banning, Magnesia Canyon, 12 mi e. Edi-
son, Jacumba), and Mexico: Baja California
Sur (San Angel).
Hoplisoides elotae R. Bohart,
NEW SPECIES
(Pigs 3, 426; 13.47)
Male holotype.—Length 7.0 mm. Black,
marked with yellow as follows: pronotal
ridge and lobe, fore- and midtibiae in front,
apical band on T-I, broad band on T-II,
T-III to VI mostly, apical band on S-II, lat-
eral spots on S-III to V; brownish-red are:
antenna in front, mediobasal spot on clyp-
eus; brown are: legs mostly, wing spots as
in Fig. 17. Pubescence inconspicuous.
Punctation moderately coarse and close on
clypeus, frons, vertex, scutum, propodeum,
mesopleuron, and metapleuron; sparse on
scutellum; punctures 1.0—2.0 PD apart on
T-L-II, closer on remaining terga; coarse
and close on S-II. Antenna relatively sim-
ple, F-I about 2X as long as broad (Fig. 3);
652 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
17. elotae S
18. subcostalis 9 14. parkeri
Figs. 1-18. Hoplisoides species. 1, Tergum I, dorsal, punctures shown in medial area only, 25x. 2—3, Head,
frontal view, 25x. 4, Flagellum, frontal view 37.5, Mesopleuron to propodeum, lateral, 18 x. 6—8, Mesopleuron
and metapleuron, lateral, 25x. 9-14, Pygidium, 30. 15-17, Forewing, 8X. 18, Forewing, 12.
clypeus 2X as broad as long; metapleural
lower pit tiny, metapleuron breadth about
1.2 MOD (Fig. 6); T-VI angled at about
70 degrees.
Female.—Length 8 mm. Flagellum more
slender, F-I about 4X as long as broad (Fig.
4), F-IX—X white in front (Fig. 4); medio-
basal clypeal spot yellow, summit of head
yellow all across; legs all brown; pronotum
reddish, reddish spots on mesopleuron and
VOLUME 99, NUMBER 4
lateroposterior propodeal area; T-I—II partly
reddish, T-III to T-VI black laterally; py-
gidial plate with moderately fine puncta-
tion, angled at about 30 degrees (Fig. 13).
Types.—Holotype ¢ (DAVIS), 8 mi se.
Elota, Sinaloa, Mexico, V-18—62 (EF D. Par-
ker). Paratypes, 7 d, 1 2, same data as ho-
lotype.
Discussion.—As indicated in the keys,
elotae is close to spendidulus and like that
species has a punctate metapleuron. How-
ever, elotae is a much browner species, and
the flagellum is stouter in both sexes.
The specific name is a noun derived from
the town of Elota.
Hoplisoides floridicus R. Bohart
Hoplisoides floridicus R. Bohart 1968:289.
Holotype ¢, Orlando, Florida (DAVIS).
A relationship to denticulatus is indicated
by the woolly S-III to V in males. However,
floridicus differs in several respects in the
male. There is no trace of a lateral denticle
on S-V, the metapleural lower pit is about
as large as the midocellus, and F-VI-VII
are nodose beneath. In both sexes the wings
are extensively brownish, and the propo-
deum (except enclosure) is coarsely and
closely punctured in posterior view. Some
females resemble those of placidus, but the
bent lateral pygidial carina of the latter is
distinctive.
I have seen 7 3 and 26 9, all from Flor-
ida.
Hoplisoides glabratus R. Bohart
Hoplisoides glabratus R. Bohart 1968:291.
Holotype ¢, Granite Pass, Hidalgo Co.,
New Mexico (DAVIS).
This little wasp is uncommonly collect-
ed. It is extensively polished and the wings
are dark except for the bright yellowish
stigma. Both sexes have the thorax and
much of the abdomen red. The male has F-I
to IV expanded beneath, but especially I
and IV. The female frons and antenna are
all or nearly all light yellow. Females fre-
653
quent the undersides of Bailey pleniradiata
flowers and emerge to snare leafhoppers.
I have studied 15 ¢ and 80 &, most
which I collected at Granite Pass, New
Mexico. Other records include Pearce, Will-
cox, Douglas, and Portal, Arizona; Deming
and San Antonio, New Mexico; Colorado
Springs, Colorado; Odessa and Marfa, Tex-
as.
Hoplisoides hamatus (Handlirsch)
Gorytes hamatus Handlirsch 1888:403. Ho-
lotype d (seen), ““Colorado” (VIENNA).
Gorytes spilographus Handlirsch 1895:895.
Holotype ¢ (seen), ‘“‘Nordamerika”’ (GE-
NEVA). Synonymy by Bohart in Bohart
and Menke 1976:521.
Hoplisoides arizonensis Baker 1907:164.
Holotype & (seen), Prescott, Arizona
(WASHINGTON). Synonymy by Bohart
in Bohart and Menke 1976:521.
Gorytes adornata Bradley 1920:115. Ho-
lotype 2 (seen), Felton, Santa Cruz Mts.,
California ITHACA). Synonymy by Bo-
hart in Bohart and Menke 1976:521.
Perhaps the most abundant species in
California, hamatus is readily recognized
by the sharply “‘beveled”’ male clypeus, the
large oval metapleural lower pit, the dis-
crete spot at the end of the subdiscoidal cell
in the female, and the all-black propodeum.
I have studied 160 d and 210 2 from
almost every county in California from sea
level to 8,000 feet in the Sierra. Out-of-state
records which suggest wide distribution in
western North America are: Garden of the
Gods, Colorado; and Palominas and Oak
Creek Canyon, Arizona.
Hoplisoides insularis (Cresson)
Harpactus insularis Cresson 1865:146. Ho-
lotype 2, Cuba (HAVANA-ID).
Characteristics are the relatively large
metapleural lower pit, generally fine punc-
tation, unbanded T-III, female F-I about
2.9X as long as greatest breadth, and sterna
as well as pygidium of female red.
I have seen only a 2 collected in Havana.
654
Alayo (1969) recorded 5 specimens from
various Cuban localities.
Hoplisoides iridipennis (F Smith)
(Fig. 10)
Gorytes iridipennis FE Smith 1856:363. Ho-
lotype ¢ (seen), Santarem, Brazil, (LON-
DON).
Gorytes fasciatipennis Cameron 1890:75.
Holotype 2 (seen), ““N. Yucatan’? (LON-
DON). Synonymy by Bohart in Bohart
and Menke 1976:521.
Gorytes maculipennis Cameron 1890:73.
Holotype ¢ (seen), Bugaba, Panama
(LONDON). New synonym.
Gorytes panamensis Maidl and Klima
1939:91. New name for maculipennis
Cameron 1890, nec Giraud 1861.
Characteristics of this wasp are: small
size (about 7 mm long (male) and 8 mm
(female)), nearly impunctate and polished
pleuron, mostly polished T-II, metapleural
lower pit oval and much larger than mid-
ocellus, T-I to V yellow banded but II to
VI sometimes mostly yellow, pygidial plate
well punctured and striatiform in part (Fig.
10), other features are: practically no sulcus
below midocellus, mandible yellow toward
base, clypeus considerably yellow, fore-
wing costa reddish and stigma yellow, scu-
tum moderately punctate, legs and venter
partly yellow, T-III with many punctures 2
to 4 PD apart.
Of the typical sort with narrowly yellow-
banded terga, I have seen 9 ¢ and 16 @
from Mexico (Vera Cruz, Oaxaca), Nica-
ragua, El Salvador, Costa Rica, Panama,
Ecuador, Venezuela, Suriname, Brazil. The
more extensively yellow form are 4 6 and
7 2 from Mexico (Vera Cruz, Sinaloa, Ja-
lisco, Morelos, Oaxaca, Puebla).
Hoplisoides jaumei (Alayo)
Psammaecius jaumei Alayo 1969:17. Ho-
lotype 2, Rangel, Pinar del Rio, Cuba
(HAVANA-1). Transferred to Hoplisoides
in Bohart and Menke 1976:521.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
I do not know this species. It is included
in the key on the basis of Alayo’s original
description of the only known specimen.
Hoplisoides jibacoa (Alayo)
Psammaecius jibacoa Alayo 1969:12. Syn-
type 6, 2, Cuba (HAVANA I).
I know this species only from the original
description, and from a male syntype sent
to me by Alayo. The male is black and lem-
on yellow, flagellum and S-III to VI all
black, and clypeus finely but closely punc-
tured. The metapleural lower pit is round
and about as large as the midocellus.
The species is known only from Cuba,
where it is moderately abundant.
Hoplisoides niger R. Bohart,
NEW SPECIES
(Pigs., 74, 14, 6)
Female holotype.—Length 8 mm. Black,
marked with yellow as follows: scape in
front, strip on lower frons next to eye, bro-
ken narrow band at apex of T-II; flagellum
dull reddish within; wing cloud as in Fig.
16, costa black, stigma brown. Pubescence
moderately appressed on clypeus, incon-
spicuous elsewhere. Punctation faint on
frons, practically absent on polished scu-
tum, pleuron, T-I—II, most of venter; quite
fine and sparse on polished T-III to V, scat-
tered and moderate on polished S-II and py-
gidial plate (Fig. 11). F-I 2.7X as long as
broad; sulcus below midocellus well devel-
oped and extending one-third of distance to
clypeus; forward part of scrobal sulcus two-
thirds complete (Fig. 7); metapleuron most-
ly 1.8X as broad as MOD, metapleural low-
er pit oval and larger than MOD (Fig. 7),
propodeal enclosure with 14 complete ca-
rinulae; T-I a little longer than broad, T-II
nearly twice as broad as long; pygidial plate
twice as long as broad, with a weak longi-
tudinal median ridge (Fig. 11).
Male.—Unknown.
Type.—Holotype 2 (WASHINGTON),
Maricao, Puerto Rico, VI-20-69 (O.S. Flint,
Jr.).
VOLUME 99, NUMBER 4
Discussion.—The practically impunctate
pleuron, large metapleural lower pit, and
complete propodeal enclosure carinulae re-
late this species to iridipennis. However,
there are many differences. Hoplisoides ni-
ger is more extensively black and even less
punctate, the midocellar sulcus is longer,
the scrobal sulcus is more nearly complete,
and the pygidial plate is quite different
(compare Figs. 10, 11).
The specific name is a noun based on the
overall black appearance.
Hoplisoides parkeri R. Bohart,
NEW SPECIES
(Figs. 1, 2, 14, 15)
Holotype male.—Length 7.5 mm. Black,
marked with yellow as follows: central spot
on clypeus, inner eye margin, apical bands
on T-I to V, VI entirely, lateral dots on S-II
to V; brownish are: legs, tegula, parategula;
forewing with clouds as in Fig. 15. Pubes-
cence inconspicuous, silvery, bloomlike,
evident and appressed on clypeus surround-
ing central spot. Punctation moderate on
head, coarse and close on thorax except
spaced 1.0 to 3.5 PD on pleuron, coarse and
0.5—1.0 PD apart on terga (Fig. 1) and S-II,
finer and more spaced on other sterna. F-I
1.5X as long as broad, F-II to X about as
broad as long, F-II to VI somewhat convex
beneath, F-VI to X with shiny spots be-
neath; metapleuron with nearly even
breadth of about 1.0 MOD, metapleural
lower pit about equal to midocellus, pro-
podeal enclosure with 12 complete longi-
tudinal carinulae, T-I slightly longer than
broad, T-VI angled at about 35 degrees.
Female.—Length 8-10 mm., clypeus
with more variegated markings and less pu-
bescent than male, yellow bands sometimes
present on pronotum and scutellum; pygid-
ium as in Fig. 14.
Types.—Holotype d (DAVIS), 4 mi nw.
Choix, Sinaloa, Mexico, VIII-31-68 (T. A.
Sears, R. C. Gardner, C. S. Glaser). Para-
types (all from Mexico): 17 ¢d, 1 2, same
data (practically) as holotype; 5 6, Puebla
(Petlalcingo), VIII-3—63, (EF D. Parker, L.
655
A. Stange); 5 6, 1 2, Hidalgo (Jacola),
VIII-31—60 (Scullen, Bolinger); 6 3, Guer-
rero (near Chilpancingo), VIII-1962 (U.
Kansas Exped.); 5 d, 1 2, Chiapas (20 mi
s. Tuxtla Gutierrez), VIII-12—63 (FE D. Par-
ker, L. A. Stange); 4 6, 2 2, Oaxaca (44
mi w. Tehuantapec), VII-21—52 (E. E. Gil-
bert, C. D. MacNeil). Also, non-paratypes,
13 6, 8 @ from various Mexican states in-
cluding Tamaulipas, Morelos, and Zacate-
cas. A few specimens from Liberia in Costa
Rica and Quezaltepeque in El Salvador are
conspecific.
Discussion.—The close and coarse punc-
tation of most of the thorax and abdomen,
relatively unmodified male antenna, fairly
large metapleural lower pit, basally black
mandible, mostly brown legs, clouding over
practically all of discoidal cell, and yellow
markings of terga increasing toward apex,
are characteristic in combination. Yellow
markings vary in both sexes. Males may
have narrow bands on the pronotum and
scutellum. T-I may or may not be all black.
Tibiae may be partly yellow.
The species is named for my friend,
Frank Parker, who collected much of the
type series.
Hoplisoides placidus (R Smith)
(Fig. 9)
Gorytes placidus F Smith 1856:368. Syn-
types d, 2 (seen), ‘East Florida” (LON-
DON).
Gorytes rufipes EF Smith 1856:369. Syntype
2 (seen), ‘‘East Florida’? (LONDON).
Synonymy by Bohart and Menke 1976:
521.
The dark wings and extensive burnt-red
coloration mark this as a typical Floridean
wasp. In general it resembles floridicus, but
the male sternal pubescence and sizable
yellow bands on T-III to V of floridicus in
both sexes are differentiating. Also, the fe-
male of placidus has the lateral pygidial ca-
rinae bent (Fig. 9). Other differences in
both sexes of placidus are the larger and
oval metapleural lower pit and the close,
656
coarse punctation of the propodeum in pos-
terior view.
I have seen 7 6 and 25 @, all from Flor-
ida.
Hoplisoides placidus nebulosus (Packard)
Gorytes nebulosus Packard 1867:424. Lec-
totype ¢d (seen), designated by Cresson
1928:48, “‘New Jersey”> (PHILADEL-
PHIA).
Gorytes armatus Provancher 1888:272. Ho-
lotype 2 (seen), Ottawa, Canada (QUE-
BEC). Synonym by Bohart in Bohart and
Menke 1976.521.
Gorytes microcephalus Handlirsch 1888:
405. Syntype d (seen), “Georgia” (GE-
NEVA). Synonymy by Bohart in Bohart
and Menke 1976:521.
Goyrtes pergandei Handlirsch 1888:407.
Syntype 6 (seen), ‘‘ Virginia and Illinois”
(GENEVA). Synonymy in Bohart and
Menke 1976:521.
Philanthus harringtonii Provancher 1888:
278. Holotype 2 (seen), Ottawa, Canada
(QUEBEC). Synonymy by Bohart in Bo-
hart and Menke 1976:521.
Gorytes birkmanni Baker 1907:166. Holo-
type @ (seen), Fedor, Texas (WASHING-
TON). New status.
Gorytes pruinosus Baker 1907:166. Holo-
type 2 (seen), Fedor, Texas (WASHING-
TON). Synonymy by Bohart in Bohart
and Menke 1976:521.
Bohart and Menke (1976) treated nebu-
losus as a subspecies of placidus, and I am
in agreement. Baker’s birkmanni is an in-
termediate form with mostly red propo-
deum but wings less dark than in typical
placidus. Males of nebulosus resemble
those of spilopterus, and characters given
in the key are not always satisfactory for
separation. However, the bent pygidial ca-
rinae of female nebulosus are distinctive.
Perhaps males are best separated by geog-
raphy, nebulosus occurring east of the
100th meridian and spilopterus west of it.
I excavated 2 ground nests of nebulosus
near Lake Texoma, Oklahoma. They were
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
provisioned with many membracid nymphs,
and a few adults.
I have studied 31 ¢ and 31 @ of this
subspecies, characterized by the large oval
metapleural lower pit, most terga with yel-
low bands, and lateral carinae of the female
pygidium (T-VI) bent, rather than evenly
curved (as in Fig. 9). Records cover most
of the United States east of the 100th me-
ridian.
Hoplisoides projectus R. Bohart
Hoplisoides projectus R. Bohart 1968:290.
Holotype ¢, Los Banos, Merced Co.,
California (SAN FRANCISCO).
This species is known only from the type
series, 4 ¢ and a 2 (DAVIS), all except
holotype from the San Joaquin Valley, Cal-
ifornia. It is related to carinatus but the
eyes are farther apart in projectus, the me-
sopleural punctation finer and more spaced,
and the female pygidium much more close-
ly and rugosely punctured.
Hoplisoides punctifrons (Cameron)
Gorytes punctifrons Cameron 1890:74. Ho-
lotype ¢d (not female) (seen), Presidio,
Texas (orig. ““Mexico”’) (LONDON).
Gorytes gulielmi Viereck 1908:408. Holo-
type 6 (not female) (seen), Bill Williams
Fork, Arizona (LAWRENCE). Synony-
my by Bohart and Menke 1976:521.
The male is somewhat like hamatus but
punctifrons has the clypeus “‘beveled”’ only
laterally, and the legs are more extensively
red. Both species have a relatively large
metapleural lower pit. The female, like ha-
matus and spilopterus, has a discrete dark
spot at the end of the subdiscoidal cell in
the forewing. However, in punctifrons, T-II
is extensively red, and T-III to V are all
black, or with remnants of bands at most.
Also, the metapleural lower pit is round
rather than oval as in spilopterus.
I have studied 67 ¢ and 60 2 of this
relatively abundant species. The distribu-
tion covers most of the United States west
of the 100th meridian. I have also seen
VOLUME 99, NUMBER 4
specimens from Mexico: Sonora (Hermo-
sillo, Magdalena) and Jalisco (Choix).
Hoplisoides pygidialis (W. Fox),
NEw STATUS
Gorytes pygidialis W. Fox 1896:528. Ho-
lotype 2 (seen), ‘““Montana’”’ (PHILA-
DELPHIA).
Bohart and Menke (1976) treated pygi-
dialis as a subspecies of costalis. | am rais-
ing it to species status since the distribu-
tions are separate, and both sexes of pygi-
dialis are readily distinguished by the ex-
tensive yellow posterad on the abdomen.
Also, female pygidialis have the pygidial
plate less narrow, rather evenly expanded.
The distribution, at least in the United
States, is west of the 100th meridian. The
27 ¢ and 33 2 studied are from: North Da-
kota (Slope Co.), Montana, Utah (St.
George, Delta), Colorado (Palisade), New
Mexico (Rodeo, Mesilla Park, Hot
Springs), Arizona (Flagstaff, Portal, Bowie,
Tucson, Grand Canyon, Chandler, Sedona,
Nogales), California (Blythe, Ripley, Bard,
Duncan, Warner Springs, Scissors Crossing,
Marinette). Mexican records are: Nayarit
(San Blas), Morelos (Alpuyaca), Hidalgo
(Pachuca), Durango (Durango), Guerrero
(Acapulco), Oaxaca (Mitla), Sinaloa (Elota,
Choix), Chiapas (Tuxtla Gutierrez), Yuca-
tan (Progreso).
Hoplisoides semipunctatus (Taschenberg)
Hoplisus semipunctatus Taschenberg 1875:
367. Holotype 2, Mendoza, Argentina
(HALLE).
It appears likely that semipunctatus is a
South American species which has been in-
troduced into southeastern U.S. presumably
by airplane, since collections have been
made near airfields. I collected females at
La Cienega, Catamarca, Argentina in 1975.
The main female characteristics are the
nearly impunctate mesopleuron, smooth
propodeal enclosure, finely punctate T-II,
metapleural lower pit about as large as the
657
midocellus, and metanotum (as well as scu-
tellum) yellow.
Of the 33 2 I have studied, 8 are from
Argentina (Rio Negro, Santa Fe, Cordova,
Catamarca), 4 are from Brazil (Nova Teu-
tonia, Catarina) and the others are from
southern U.S. as follows: 4 from Alabama
(Decator), 1 from Louisiana (Baton Rouge),
1 from South Carolina (Columbia), 1 from
Mississippi (Gulfport), and 14 from Florida
(Quincy, Escambia Co., Archbold Reserve).
Hoplisoides spilopterus (Handlirsch)
Gorytes spilopterus Handlirsch 1888:414.
Syntype 2 (seen), “Nevada” (VIENNA).
Gorytes pogonodes Bradley 1920:114. Ho-
lotype 6 (seen), Lemon Cove, Tulare
Co., California ITHACA). Synonymy by
Bohart and Menke 1976:521.
This abundant species in western U.S. is
recognized by the relatively large and oval
metapleural lower pit, the subdiscoidal cell
in the female with a discrete apical spot, the
mostly red propodeum (except enclosure) in
the female, T-I to IV (at least) pale banded,
male clypeus “‘beveled’”’ but not sharply,
and clear membrane of submarginal cell I,
discoidal cell I, and medial cell. Males in
California have the legs mostly black and
yellow. Those from other western states
usually have them partly red.
I have studied 70 6 and 90 2. The dis-
tribution is widespread in California at low
to moderate altitudes. Records are also from
Arizona, Idaho, Nevada, Utah, Wyoming,
Colorado, and New Mexico, all west of the
100th meridian.
Hoplisoides splendidulus (Bradley)
Gorytes splendidula Bradley 1920:113. Ho-
lotype ¢ (seen), Brawley, California.
(ITHACA).
This elegant species, with its long, slen-
der antennae, punctured metapleuron,
sharply delineated wing clouds (including a
definite one in both sexes apically in the
subdiscoidal cell), bloomlike pubescence on
T-V, and small metapleural lower pit, are
658
all characteristic of the similar species, elo-
tae. There are a number of color differences
such as in splendidulus the much more ex-
tensive red coloration, the all dark red
T-IV—V, and the red (or yellow) frons.
More important is the more slender flagel-
lum in splendidulus. For example, F-IV is
1.8X (male) to 2.8X (female) as long as
broad. In elotae, F-IV is 1.4 (male) to
1.8X (female) as long as broad.
I have studied 13 6 and 11 @ from Texas
(Starr Co., Ward Co.), New Mexico (Las
Cruces), Arizona (Tucson, Prescott, Grand
Canyon, Yuma Co.), Utah (Nephi), Oregon
(Antelope Mt., Harney Co.), California
(Jacumba, Warner Springs, Brawley, La-
guna Canyon, Antelope Springs, Johnsville,
Tesla, Napa Co., San Diego Co., Blythe),
and Mexico: Baja California Sur (San Vin-
cente).
Hoplisoides subcostalis R. Bohart,
NEW SPECIES
(Figs. 5, 127 18)
Holotype female.—Length 10 mm.
Black, with yellow markings as follows: F-I
to V within, scape in front, mandible to-
ward base, clypeus partly, frons laterally,
narrow band on pronotal ridge, lobe partly,
narrow posterior band on scutellum, fore-
and midtibiae and tarsi partly, faint apices
of T-II to IV; reddish brown are: flagellum
toward apex, legs mostly; pygidium dark
red; forewing black across entire front (Fig.
18). Pubescence inconspicuous. Punctation
practically absent on frons, moderate and
1-2 PD apart on scutum, mostly fine and
widely spaced on mesopleuron, absent to-
ward base of propodeal side, moderately
coarse and close on propodeum posteriorly,
fine and widely spaced on T-I-II, increas-
ingly closer on T-III to V, coarse and mod-
erately close on pygidium. Mandible gently
curved and moderately stout (as compared
with stouter one of costalis), posterior meta-
pleural margin undefined but followed by a
series of diagonal carinae at propodeal base
(Fig. 5). Metapleural lower pit tiny; pro-
podeal enclosure completely carinulate; for-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ebasitarsus posteriorly black, T-I about as
long as broad, T-II about 1.3 as broad as
long, pygidial plate narrow overall but ex-
panding evenly above (Fig. 12).
Male.—Unknown.
Types.—Holotype 2 (WASHINGTON),
Barro Colorado, Canal Zone, Panama, III-
15-67 (R. D. Akre). Paratype, 2 (DAVIS),
topotype, IV-6—63 (C. & M. Rettenmeyer).
Discussion.—This species is related to
costalis on the characters of the clypeal
conformation, relatively large size, small
metapleural lower pit, and narrow pygidial
plate. On the other hand, it differs by the
finer punctation of the pleuron, T-I—II, and
the frons, but especially by the black ante-
rior one-third of the forewing. The more
slender mandible in subcostalis is also dif-
ferent.
Hoplisoides tricolor (Cresson)
(Fig. 8)
Gorytes tricolor Cresson 1868:380. Holo-
type £ (seen), New Mexico (PHILA-
DELPHIA).
Gorytes helianthi Rohwer 1911:569. Holo-
type 2 (seen), Boulder, Colorado
(WASHINGTON). Synonymy in Bohart
and Menke 1976:521.
Hoplisus rufocaudatus Mickel 1916:401.
Holotype 2 (seen), Mitchel, Nebraska
(LINCOLN). Synonymy in Bohart and
Menke 1976:521.
This species is characterized by the ex-
tensively red markings, especially on the
legs, propodeum, and T-I (rare exceptions).
Otherwise, the small lower pit at the bottom
of a tapering metapleuron (Fig. 8), dark-red
last two or three terga, usually exposed
T-VII in the male, extensively clouded sub-
discoidal cell, and broadly yellow female
frons are more critical characters.
I have studied 53 6 and 27 2 from Texas
(Llano Co.), Wyoming (Grand Teton Park),
Colorado (Palisades, Hasty, Pueblo), New
Mexico (Lordsburg, White Sands Monu-
ment, Rodeo, Correo, Granite Gap, Tu-
cumcari, near Deming), Utah (Delta, Cor-
VOLUME 99, NUMBER 4
nish, Salt Lake City, W. Utah Lake, Logan),
Arizona (Toltec, Eloy, Willcox, Portal, An-
imas, Tucson, Huachuca Mts.), California
(Westmorland, Tracy, Warner Springs, Eliz-
abeth Canyon in Los Angeles Co.), Mexico:
Sonora (Alamos), Sinaloa (Mazatlan,
Choix), Nueva Leon (Apodaca), Coahuila
(Saltillo). Except for Llano Co., Texas
(99°), all of the United States localities are
west of the 100th meridian.
Hoplisoides vespoides (F. Smith)
Gorytes vespoides F Smith 1873:407. Ho-
lotype ¢ (seen), Ega (now Tefé), Brazil
(LONDON).
Gorytes robustus Handlirsch 1888:380.
Syntype ¢ (seen), Blumenau, Brazil (VI-
ENNA); Tampico, Mexico (GENEVA).
Synonymy by Bohart in Bohart and Men-
Kkep'976:5 21%
Icuma sericea Cameron 1905:21. Syntype
2 (seen), ‘““Panama’’ (LONDON). Syn-
onymy by Bohart in Bohart and Menke
19762521:
Gorytes auropilosellus Cameron 1912:430.
Holotype 2 (seen), “British Guiana”’
(LONDON). Synonym by Bohart in Bo-
hart and Menke 1976:521.
Hoplisoides umbonicida Pate 1941:1. Ho-
lotype @ (seen), Caura Valley, Trinidad.
(PHILADELPHIA). Callan (1976:332)
suggested the synonymy. New synonym.
This is one of the largest species of Hop-
lisoides, females often with length of 12—
13 mm. The females are robust, and most
of them have T-I dark, T-II to V with apical
yellow bands. Species characteristics are
the relatively large oval metapleural lower
pit, weak and isolated mesopleural punc-
tures, extensive reddish brown wing color-
ation, becoming darker apically, T-II about
2X as broad as long, pygidial plate of fe-
male nearly 2X as long as broad, and pro-
podeal enclosure with at least 20 carinulae
but without distinct lateral boundaries.
I have studied 7 d and 15 ¢ from Mex-
ico: Chiapas (Tuxtla Gutierrez), Nayarit
(near Tepic), Morelos (Alpuyeca, Cuerna-
659
vaca, Lake Tequesquitengo), Guerrero
(Chilpancingo), Yucatan (Chichen Itza).
Specimens seen from other countries are:
Guatemala (Lake Amatitlan), El Salvador
(Los Charros), Panama (Potrerillos, Frejo-
les Canal), Ecuador (Azuay Prov., Limon-
cocha), Trinidad (Mundo Nuevo), Suriname
(Paramaribo), Peru (Colonia Ferane), Brazil
(Obidos in Para, Itatiaya, Nova Teutonia).
Two @ in the collection are pinned with
their prey, membracid adults, which must
outweigh them.
Hoplisoides xerophilus Alayo
Psammaecius confusus Alayo 1969:14.
Syntype 6, 2, Cuba (HAVANA JI).
Preocc. by Dutt 1922.
Hoplisoides xerophilus Alayo 1976:29.
New name for confusus Alayo.
This species is known to me only by a
male syntype sent by Alayo. The relatively
large metapleural lower pit, completely
carinulate propodeal enclosure, and un-
banded T-III are distinctive in combination.
The moderately punctured pleuron and T-II,
particularly in the female as pictured by
Alayo (1969), are additional characters. Ac-
cording to Alayo (1969), the species is fair-
ly common in coastal localities of Cuba at
flowers of Coccolaba unifera.
LITERATURE CITED
Alayo, D. P. 1969. Studios sobre los Himenopteros de
Cuba. HI. Subfamilia Nyssoninae. Poeyana (ser.
A). No. 59: 1-34.
. 1976. Introduccién al estudio de los Hime-
ndépteros de Cuba. Academia de Ciencias de Cuba,
Serie Biologia, No. 67: 2—46.
Baker, C. E 1907. Some new Gorytes-like wasps. In-
vertebrata Pacifica 1: 161-178.
Bohart, R. M. 1968. New Hoplisoides from the Unit-
ed States. Proceedings of the Entomological So-
ciety of Washington 70: 287-292.
Bohart, R. M. and A. S. Menke. 1976. Sphecid Wasps
of the World. A Generic Revision. University of
California Press, Berkeley. vii-ix + 695 pp.
Bradley, J. C. 1920. Descriptions, records and notes
on North American Nyssonidae. Transactions of
the American Entomological Society 46: 113—
1322
Callan, E. McC.
1976. Observations on the nesting
660
behavior and prey of gorytine wasps in Trinidad.
Psyche 83: 324-335.
Cameron, P. 1890. Insecta Hymenoptera, vol. 2 pp.
65-128 (Fossores), xi + 413 pp. Jn Godman, F
O. and D. Salvin, eds., Biologica Centrali-Amer-
icana. Taylor and Francis, London.
. 1905. A new genus and species of Larridae
from Central America. Entomologist 38: 21—22.
. 1912. The Hymenoptera of the Georgetown
Museum. Pt. IV. The fossorial Hymenoptera. Ti-
mehri (3)2(2): 413—440.
Cresson, E. T. 1865. On the Hymenoptera of Cuba.
Proceedings of the Entomological Society of Phil-
adelphia 4: 1—425.
1868. Catalog of a small collection of Hy-
menoptera made in New Mexico during the sum-
mer of 1867. Transactions of the American En-
tomological Society 1868: 375-388.
. 1872. Hymenoptera Texana. Transactions of
the American Entomological Society 4: 153-285.
1928. The types of Hymenoptera in the
Academy of Natural Sciences of Philadelphia oth-
er than those of Ezra T. Cresson. Memoirs of the
Entomological Society 5: 1—90.
Dahlbom, A. G. 1842. Dispositio methodica speci-
erum Scandinavicarum ad familias Hymenopte-
rorum naturales pertinentium. C. Berling, Lund.
16 pp.
. 1845. Hymenoptera Europaea praecipue bo-
realia etc., fasc. 3, i—xlivt pp. 353-528.
Dalla, Torre, C. G. de. 1897. Catalogus Hymenopte-
rorum, vol. 8 (Fossores) G. Engelmann, Lipsiae.
vill + 749 pp.
Fabricius, J. C. 1775. Systema entomologiae, etc.
Kortii, Flensburgi, et Lipsiae. xxviii + 832 pp.
. 1794. Entomologia systematica emendata et
aucta. Tome IV, Hafniae. 472 pp.
Fox, W. J. 1893. New species of fossorial Hymenop-
tera. Canadian Entomologist 25: 113-117.
. 1896. Synopsis of the North American spe-
cies of Gorytes Latr., part 3. Proceedings of the
Academy of Natural Sciences of Philadelphia 46:
517-539.
Gmelin, J. E 1790. Caroli a Linné. Systema naturae
per Regnum Animale, part 5, pp. 2250—3020, G.
E. Beer, Lipsiae.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Handlirsch, A. 1888. Monographie der mit Nysson
und Bembex verwandten Grabwespen. Sitzung-
berichte Academia Wissenschaften, Wien Math-
Nat. Classe 97: 316-565.
. 1895. Monographie der mit Nysson und Bem-
bex verwandten Grabwespen. Sitzungberichte Ac-
ademia Wissenschaften, Wien Math-Nat. Classe
104: 801-1079.
Maidl, E and A. Klima. 1939. In Hedicke, Hyme-
nopterorum Catalogus 8: 3-150.
Mickel, C. E. 1916. New species of Hymenoptera of
the superfamily Sphecoidea. Transactions of the
American Entomological Society 42: 399—434.
Packard, A. S. 1867. Revision of the fossorial Hy-
menoptera of North America I. Crabronidae and
Nyssonidae. Proceedings of the Entomological
Society of Philadelphia 4: 1—425.
Pate, V. S. L. 1941. Two new species of sphecid
wasps from Trinidad. Notulae Naturae 91: 1-8.
1947. On the gorytine wasps of the West
Indies. Entomological News 58: 93-98.
Provancher, |’Abbé L. 1888. Additions et corrections
au volume II de la Faune Entomologique du Can-
ada traitant des Hyménopteres. pp. 1—438.
Rohwer, S. A. 1911. Descriptions of new species of
wasps with notes on described species. Proceed-
ings of the United States National Museum 40:
551-587.
Smith, EF 1856. Catalogue of hymenopterous insects
in the collection of the British Museum, part IV,
Sphegidae, Larridae, and Crabronidae, pp. 207—
497. London.
. 1873. Descriptions of new species of fosso-
rial Hymenoptera in the collection of the British
Museum. Annals and Magazine of Natural His-
tory (4)12: 402-415.
Taschenberg, E. 1875. Nyssonidae und Crabronidae
des Zoologisches Museums der hiesigen Univer-
sitaét. Zeitschrift fiir Naturwissenschaften. Halle
(new series 2) 45: 359-409.
van der Vecht, J. 1961. Hymenoptera Sphecoidae Fa-
briciana. Zoologische Verhandelingen 48: 3-85.
Viereck, H. L. 1908. Notes and descriptions of Hy-
menoptera from the western United States. II. Bo-
real species. Transactions of the American Ento-
mological Society 33: 381—408.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 661—665
LESTES JERRELLI, N. SP. (ZYGOPTERA: LESTIDAE),
A NEW DAMSELFLY FROM ECUADOR
K. J. TENNESSEN
1949 Hickory Ave., Florence, AL 35630, U.S.A.
Abstract.—Lestes jerrelli, n. sp., is described and illustrated from 13 males and 8
females (holotype d, allotype 2, in copula: Ecuador, Napo Prov., pond 12.3 km W of
Coca, elev. 250 m, 13 June 1995). It is related to L. jurzitzai Muzon and L. paulistus
Calvert from Brazil, but is distinct in thoracic color pattern and shape of male paraprocts.
Key Words:
Lestes is a relatively large genus of dam-
selflies, with about 80 species known
worldwide (Bridges 1993), nearly half of
these occurring in the western hemisphere.
The latest description of a new species from
tropical America was that of Lestes jurzitzai
Muzon (1994) from Brazil. The new spe-
cies described here was collected east of the
Andes Mountains in Napo Province, Ecua-
dor. It is mostly blue with narrow metallic
green thoracic stripes, and is most closely
related to Lestes jurzitzai based on color
pattern and male appendage morphology.
The new species was collected at a small
pond and several small, shallow, temporary
pools west of Coca, near the equator
(0°27'S, 77°0'W). Aquatic vegetation var-
ied from sparse to abundant.
Lestes jerrelli Tennessen, NEW SPECIES
(Figs. 1-4, 8)
Holotype 6: ECUADOR, Napo Prov-
ince, pond along Loreto Road, 12.3 km W
of junction with Coca Road, elev. 250 m,
13 June 1995, W. M. Mauffray, leg.; de-
posited in Florida State Collection of Ar-
thropods (FSCA), Gainesville, Florida,
USA.
Allotype 2: in copula with holotype; de-
posited with holotype.
Damselfly, Lestes jerrelli, new species, Ecuador
Paratypes (12 36, 7 2): Same data as ho-
lotype, 3 tandem pairs, K. J. Tennessen leg.
(Coll. K. J. Tennessen); same data, 3 tan-
dem pairs, W. M. Mauffray leg. (FSCA, In-
ternational Odonatological Research Insti-
tute, Catholic Univ., Quito). ECUADOR,
Napo Prov., temporary pool along Loreto
Rd., 1.7 km W of junction with Coca Rd.,
13 June 1995, K. J. Tennessen leg., 4 d, 1
tandem pair (KJT, Coll. R. W. Garrison);
same data, 1 ¢6, W. M. Mauffray leg., (Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C.).
Other specimens (not examined): In ad-
dition to Ecuador, Lestes jerrelli occurs in
southern Peru: 2 d in the Collection of R.
W. Garrison and 2 ¢ and 1 & in the Na-
tional Museum of Natural History, Smith-
sonian Institution, are from Madre de Dios
Dept., Hotel Amazonia, across from Ata-
laya, collected 27 June 1993.
Etymology.—This species is named for
Jerrell J. Daigle in recognition of his con-
tributions to New World odonatology and
his enthusiasm for the study of dragonflies.
Holotype male.—General color pattern
light blue with dark green metallic mark-
ings; eyes in life medium blue dorsally,
light blue ventrally.
Head: Tips of mouthparts black; labrum
662
Figs. 1-3.
3, Male appendages, lateral view.
sky blue, distal margin black, and with a
small, triangular, mediobasal black spot;
base of mandible and gena blue, a lighter
hue than labrum; anteclypeus mostly black,
partly blue laterally; postclypeus, antefrons,
antenna and postfrons black; occiput dark
metallic green, with a small, blue tear-
shaped postocular spot posterolateral to
each lateral ocellus; rear of head greenish
blue on upper half, tan near occipital fora-
men, black on lower half, partly pruinose.
Thorax: Prothorax largely blue dorsally,
but middle lobe with a submedian pair of
dark brown, outwardly curved stripes bare-
ly extending to anterior and posterior lobes,
ending in small, oblong spots; epimeron |
dark brown above, blue below. Pterothorax
largely sky blue, except as follows: mid-
dorsal stripe tan blue (light blue in life),
about 0.3 mm wide at midlength, narrowed
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Lestes jerrelli, male paratype. 1, Venter of male pterothorax. 2, Male appendages, dorsal view.
at each end by adjacent metallic, dark-green
dorsal (mesepisternal) stripes 0.3 mm wide,
parallel sided; katepisternum 2 with upper
dark brown spot; mesopleural (humeral) su-
ture light brown, posterior to which is an
isolated, metallic, dark-green stripe about
0.3 mm wide, constricted in posterior
fourth; metepimeron with an anterior, black
oval spot visible in lateral view and extend-
ing anteromedially as a thin line nearly to
midline of thoracic venter, and with a small-
er posterior black spot below posterior ca-
rina, also visible in lateral view (Fig. 1);
metapostepimeron blue; thoracic venter
pale tan, pruinose, and with a diffuse, nar-
row, dark lateral stripe on each side and
along the invaginated midline anterior to
poststernum. Coxae partly pruinose, pro-
and mesocoxae blue and light brown, meta-
coxa all brown; profemur black with inner
VOLUME 99, NUMBER 4
Figs. 4-7.
view. 4, Lestes jerrelli, Peru, Madre de Dios Dept. 5,
Lestes pictus, Brazil, Santa Catarina State. 6, Lestes
jurzitzai, Brazil, Rond6énia State. 7, Lestes paulistus,
Brazil, Santa Catarina State. (Figs. 4—7 drawn by R.
W. Garrison.)
Left male cercus and paraproct, dorsal
surface pale tan in basal half, meso- and
metafemora mostly black but with tan
stripes on dorsal and ventral surfaces; tibiae
black except extensor surfaces tannish blue;
tarsi and claws black. All tergal and axillary
sclerites blue; wings hyaline, with 13 or 14
postnodal crossveins in fore wing, 12 in
hind wing; pterostigma dark brown, sur-
mounting 2 cells, 1.40 mm long in hind
wing.
Abdomen: (Abdominal segments desig-
nated with an S and a number, e.g., Sl =
segment 1). S1 blue with very small sub-
median and sublateral dark spots, postero-
lateral and ventral carinae black; S2 blue
with elongate dorsolateral dark metallic
663
8
Figs. 8-9. Posterior margin of male abdominal
segment 10, Dorsolateral view. 8, Lestes jerrelli, Ec-
uador, Napo Prov. 9, Lestes jurzitzai, Brazil, Rond6nia
State. (Fig. 9 drawn by R. W. Garrison.)
green stripe about 0.4 mm wide, not reach-
ing posterior margin of segment; S3 about
2.3 times as long as S2; S3 to S7 largely
dark metallic green, S3 with narrow blue
basal ring, S4 to S7 with small, blue, ba-
solateral triangles, these connecting to ven-
trolateral blue stripes anteriorly and tan
stripes posteriorly; S8 with narrow, dark bas-
al ring, S8 and S9 pruinose, dark brown
dorsally, reddish brown laterally; posterior
margin of S10 with dorsal, stout, black
spines (Figs. 2, 8). Cercus black, about 1.5
times longer than S10, in lateral view de-
curved in apical fourth (Fig. 3), in dorsal
view curved inwardly (Fig. 2), outer margin
with numerous stout spines, elongate dorsal
subapical concavity present, basal tooth
sharp, quadrate, ventromedial lobe slightly
convex with about 6 to 8 small marginal
teeth and a few smaller denticles; para-
procts black, in lateral view straight (Fig.
3), in dorsal view relatively stout (0.14 mm
wide at narrowest point), directed inward
and expanded at tip, dorsal surface of tip
with shallow depression (Fig. 2).
Measurements (mm): Total length (in-
cluding cercus) 40.0, abdomen length 32.0,
hind wing length 21.2.
Allotype female:—Color pattern similar
to male, blue hue less intense; other differ-
ences noted below.
Head: Labrum without mediobasal black
spot; postclypeus dark brown; base of an-
tenna and segments | and 2 partly tan; me-
dian ocellus surrounded by narrow tan
664
marking, lateral ocelli with oblong tan spot
laterally.
Thorax: Prothorax greenish blue; ptero-
thorax with blue area more extensively in-
vaded by tan than in male; legs with pale
stripes wider and longer than in male;
wings with 13 postnodal crossveins in fore
wing, 12 in hind wing; pterostigma 1.50
mm long.
Abdomen: S7 about 2.5 times as long as
S8; apical segments darker than in holo-
type, S8 with dorsomedial, light brown oval
spot; posterolateral corner of basal plate of
Ovipositor truncate; gonostyle 0.48 mm
long; tips of ventral valves extending to
posterior margin of S10; cercus 0.60 mm
long, about 3/4 length of S10.
Measurements (mm): Total length (in-
cluding cercus) 37.3, abdomen length 29.2,
hind wing 21.7.
Variation in paratypes.—Male: labrum
sky to medium blue, mediobasal black spot
oval or triangular; katepisternum sometimes
entirely blue; post-mesopleural metallic
green stripe sometimes interrupted in apical
fourth; wings sometimes slightly flavescent;
12 to 15 postnodals in fore wing, 11 to 15
in hind wing; pterostigma 1.15—1.40 mm
long; Sl sometimes without small black
spots. Total length 38.0—41.0 mm; abdo-
men length 30.5—33.0 mm; hind wing
length 19.6—22.3 mm. Female: labrum
sometimes with mediobasal black spot;
wings usually flavescent; 12 to 14 post-
nodals in fore wing, 11 to 14 in hind wing;
pterostigma 1.40—1.55 mm long; postero-
lateral corner of basal plate of ovipositor
sometimes acutely angulate; cercus 0.56—
0.62 mm long. Total length 36.0—39.5 mm;
abdomen length 28.0—30.7 mm; hind wing
length 21.0—23.3 mm.
COMPARISON WITH OTHER SPECIES
Lestes jerrelli belongs to a small group
of South American species that have bright
blue markings which includes L. jurzitzai
and L. paulistus Calvert (1909) from Brazil,
L. pictus Hagen in Selys (1862) from Brazil
and Argentina, and L. debellardi De-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Marmels (1992) from Venezuela. In L. pic-
tus and L. debellardi, the male cercus lacks
a basal tooth (Fig. 5), and the paraproct is
only about half as long as the cercus. Lestes
jerrelli differs from L. jurzitzai and L. pau-
listus by the tips of the paraprocts being
expanded and dorsally excavated (Figs. 2,
4); in most specimens of L. jerrelli, the tip
of one of the paraprocts overlies the other.
In dorsal view, the paraprocts of L. jurzitzai
are parallel-sided and curve outward slight-
ly (Fig. 6), whereas those of L. paulistus
are acuminate, shorter, and angled inward
(Fig. 7). The dorsal, posterior margin of ab-
dominal segment 10 bears larger, stout
spines in L. jerrelli (Fig. 8) compared to L.
jurzitzai (Fig. 9). The dark thoracic stripes
and anterior abdominal markings of L. jur-
zitzai are black (Muzon 1994); these mark-
ings in L. paulistus were black in specimens
examined by Muzon (1994), although a
male in the Collection of R. W. Garrison
has metallic green thoracic stripes (personal
communication). All specimens of L. jer-
relli | examined have dark-green metallic
thoracic stripes. In L. jerrelli, the dark-
green stripe posterior to the mesopleural su-
ture is complete at its anterior end, whereas
in L. jurzitzai this stripe is interrupted an-
teriorly (see Muzon 1994, Fig. 1a); also in
L. jurzitzai, the mesepisternal dark stripe
has a lateroventrally directed arm at its pos-
terior end. The color pattern on the venter
of the pterothorax in L. jerrelli is similar to
L. jurzitzai and L. pictus (see Calvert 1909,
Plate 1, Fig. 11). However, Calvert’s figure
of pictus shows that the metepimeral black
spot in L. pictus is not elongated into a nar-
row, anteromedially-directed stripe as in L.
jurzitzai (see Muzon 1994, Fig. 1B and L.
Jerrelli, Fig. 1).
ACKNOWLEDGMENTS
I thank Bill Mauffray for the loan of
specimens. I am grateful to Rosser Garrison
for valuable comments on the manuscript
and comparative notes on all four species,
and for providing drawings for Figs. 4—7
VOLUME 99, NUMBER 4
and Fig. 9, and Thomas W. Donnelly for
reviewing the manuscript.
LITERATURE CITED
Bridges, C. A. 1993. Catalogue of the Family-group,
Genus-group and Species-group Names of the
Odonata of the World, 2nd ed. Charles A. Bridges,
Urbana, Illinois, 806 pp.
Calvert, P PR 1909. Contributions to a knowledge of
the Odonata of the neotropical region, exclusive
665
of Mexico and Central America. Annals of the
Carnegie Museum 6(1): 73-281.
DeMarmels, J. 1992. Dragonflies (Odonata) from the
Sierras of Tapirapeco and Unturan, in the extreme
south of Venezuela. Acta Biologica Venezuelica
14(1): 57-78.
Muzon, J. 1994. Lestes jurzitzai spec. nov., a new
damselfly from Rond6nia, Brazil (Zygoptera: Les-
tidae). Odonatologica 23(3): 297-302.
Selys, M. E. 1862. Synopsis des Agrionines. Seconde
légion: Lestes. Bulletin de 1’ Académie Royale de
Belgique (2)13: 288-338.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 666-675
SYNONYMY OF TWO GENERA (EUMAYRIA AND TRISOLENIELLA) OF
CYNIPID GALL WASPS AND DESCRIPTION OF A NEW GENUS,
EUMAYRIELLA (HYMENOPTERA:CYNIPIDAE)
GEORGE MELIKA AND WARREN G. ABRAHAMSON
Department of Biology, Bucknell University, Lewisburg, PA 17837, U.S.A.; (GM) per-
manent address: Savaria Museum, Kisfaludy S.U. 9, Szombathely, 9701, Hungary.
Abstract.—The genus Trisoleniella Rohwer and Fagan is a new synonym of Eumayria
Ashmead. A redescription of Eumayria and a key to species are given. Species included
in Eumayria are: E. bignelli (Dalla Torre and Kieffer), n. comb.; E. brevicornis (Beuten-
mueller), n. comb.; E. enigma (Weld), n. comb.; E. floridana Ashmead; and E. saltata
(Ashmead), n. comb. Eumayria enigma is probably the unisexual generation of E. flori-
dana. Two species previously placed in Eumayria are transferred to other genera: E.
longipennis (Ashmead) to Andricus (n. comb.) and E. invisa (Weld) to Eumayriella (n.
comb.). Eumayriella, n. gen., is described to include E. archboldi, n. sp., and E. invisa,
and it is compared to the closely related genus Eumayria.
Key Words:
The diagnostic features currently used in
the keys to North American oak cynipid
genera of the tribe Cynipini frequently in-
clude morphological characters that are
variable and consequently are unable to dis-
tinguish different genera. These systematic
difficulties are the outcome of using inap-
propriate diagnostic characters for some
genera. The presence or absence of a tooth
of the tarsal claws is the main diagnostic
character used to divide all the Cynipini
into two large generic groups and the cur-
rent systematic arrangement of the tribe is
based primarily on this characters (Weld
1952a). Ashmead (1886), in his earlier key
to the genera of the Cynipidae did not use
this criterion; however, his later generic key
included this characteristic (Ashmead
1903). This morphological criterion is in-
sufficient for all taxonomic distinctions.
The presence or absence of tooth on the tar-
sal claw is likely a homoplasy and probably
evolved independently in several Cynipini
cynipoids, gall wasps, taxonomy, morphology, distribution, biology
genera. The ancestral condition is probably
the simple tarsal claw since it appears in the
majority of genera from the Aylacini. The
authors will analyse this character in detail
in another work.
Furthermore, the nomenclature used to
describe taxonomically important structures
in publications prior to the 1960s is incon-
sistent (Dailey and Menke 1980). Various
authors have used different terminology to
define the same morphological structure
and have employed the same term for dif-
ferent structures. As a consequence, species
have been misunderstood and misidentified.
Four species of Eumayria Ashmead were
known, all from the United States: E. el-
doradensis (Beutenmueller), E. floridana
Ashmead, E. invisa Weld, and E. longipen-
nis (Ashmead). Eumayria eldoradensis was
transferred into Callirhytis previously
(Dailey et al. 1974), while E. longipennis
herein is transferred to Andricus. A com-
parison of the morphological criteria des-
VOLUME 99, NUMBER 4
ignated earlier as diagnostic for the sepa-
ration of Eumayria and Trisoleniella and an
analysis of the types of Eumayria and Tri-
soleniella species, indicate that the two gen-
era are synonyms. Trisoleniella included
the unisexual generations, while Eumayria
contained the bisexual ones. Weld (1952a)
wrote that Trisoleniella ‘“‘may prove to be
asexual form of Eumayria’’. Consequently,
the four known species of Trisoleniella:
brevicornis (Beutenmueller), enigma
(Weld), bignelli (Dalla Torre and Kieffer),
and saltata (Ashmead) are transferred to
Eumayria. Furthermore, an examination of
the morphological characteristics of the
only known brachypterous species, E. in-
visa, showed marked differences from those
of E. floridana as well as other known Tri-
soleniella species. These differences indi-
cate that E. invisa and E. archboldi, (new
species), must be separated into a new ge-
nus Eumayriella.
We follow the current terminology for
morphological structures (Eady and Quin-
lan 1963, Fergusson 1995, Gibson 1985,
Menke 1993, Ritchie and Peters 1981, Ron-
quist and Nordlander 1989). The term “‘tho-
rax’’ used here includes the propodeum and
thus is equivalent to the ““mesosoma”’ or
‘‘mesosoma+metasoma”’ of American lit-
erature. Abbreviations for fore wing vena-
tion follow Ronquist and Nordlander
(1989). Measurements and abbreviations
used here include: FI—F12, Ist and conse-
quent flagellomeres; POL (post-ocellar dis-
tance) is the distance between the inner
margins of the posterior ocelli; OOL (ocel-
locular distance) is the distance from the
outer edge of a posterior ocellus to the inner
margin of the compound eye.
Eumayria Ashmead
Eumayria Ashmead 1887: 147; Ashmead
1903: 153. Type species: Eumayria flor-
idana Ashmead, by monotypy.
Trisolenia Ashmead 1887: 142: Ashmead
1903: 155; Dalla Torre and Kieffer 1910:
453 (as synonym of Andricus Hartig).
Type species: Andricus saltatus Ash-
667
mead, by monotypy. Preoccupied by Tri-
solenia Ehrenberg 1861 in Protozoa.
New synonymy.
Trisoleniella Rohwer and Fagan 1917: 377.
New name for Trisolenia Ashmead.
Comments.—The original descriptions of
Eumayria and Trisoleniella were made by
Ashmead (1887). However, these descrip-
tions lack sufficient precision and detail and
do not include some important diagnostic
characters. Some of the characters used in
Ashmead’s generic descriptions are found
in other genera as well. Too much attention
was paid to the description of antennae of
males and females while important charac-
ters of the head and thorax were not men-
tioned. Finally, a redescription of the genus
is necessary in order to set new limits for
the genus because of synonymization of
Trisoleniella to Eumayria.
Diagnosis.—Head 1.5—1.7 times broader
than long from above in bisexual generation
and 2.5—2.8 times broader than long in uni-
sexual females; malar groove absent. Tho-
rax flattened dorsoventrally, not arching in
anterior part; scutum slightly longer than
broad, finely coriaceous or macroscopically
punctate, never transversely sculptured.
Base of 2nd abdominal tergite with pale
felt-like ring of dense short setae, inter-
rupted dorsally (in males dense short pale
setae present only ventrally and ventroater-
ally). See also the diagnostic criteria of Eu-
mayriella below.
Redescription.—Female. Head from
above equal or broader than thorax; 1.5—1.7
times broader than long from above in bi-
sexual females (Fig. 1) and 2.5—2.8 times
broader than long in unisexual females (Fig.
6); in front view head 1.2—1.3 times broader
than high. Gena broadened behind eye,
equal or broader than cross diameter of eye.
POL and OOL nearly equal. Occiput and
vertex finely coriaceous, while frons and
gena dull rugose, frons with uniformly dis-
tributed short white setae. Malar space
slightly shorter than length of eye, without
groove, but with numerous striae radiating
668 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
e)
O
co)
ni}
Ly,
an
SSN
Figs. 1-12. 1-5, Eumayria floridana. 1, Head from above. 2, Head in front view. 3, Gaster of female in
profile. 4, Thorax in dorsal view. 5, Male antenna. 6-9, E. enigma. 6, Head from above. 7, Head in front view.
8, Gaster of female in profile. 9, Antenna. 10-12, Eumayriella archboldi. 10, Head in front view. 11, Antenna.
12, Thorax in dorsal view.
VOLUME 99, NUMBER 4
from mouthparts into malar space and frons
(Figs. 2, 7); striae more distinct and thicker
in former Trisoleniella than in Eumayria
floridana. Antenna 14-segmented in bisex-
ual females and 15—16-segmented in uni-
sexual females; in some E. floridana spec-
imens, suture separates 15th segment. F1 as
long as pedicel, scape as broad as long; F2
to F4 slightly longer than broad, subsequent
flagellomeres, except last one, subequal,
nearly as broad as long (Fig. 9). Pronotum
in median dorsal line longer than usual,
with two very dense patches of white short
setae on both sides of median dorsal line,
collar of pronotum not emarginated, lie in
same range as scutum, much broader than
usual for Cynipini (Fig. 4), and longitudi-
nally rugose. Scutum bare, very slightly
longer than broad, finely alutaceous or mac-
roscopically punctate, never transversely
sculptured; notauli usually reaching prono-
tum; parapsidal, anterior parallel and me-
dian lines present, latter distinct at least
over posterior two thirds. Scutellum bare,
equal or very slightly longer than broad,
rounded posteriorly, disk dull rugose; fo-
veae deep, with shiny, more finely sculp-
tured bottom. Mesepisternum longitudinally
rugose, posterior third smooth, shiny, with
very finely rugose sculpture. Propodeum
with smooth, bare and shiny median portion
limited by distinct carinae, converging
slightly outwards in the middle. Sides of
propodeum bearing dense white setae. Fore
wing margins without cilia, veins thick,
dark brown, areolet indistinct, very small,
triangular. Base of 2nd abdominal tergite
with pale felt-like ring of dense short setae,
interrupted dorsally. In E. floridana gaster
longer than high, 2nd abdominal tergite oc-
cupies at least % of gaster length in dorsal
view; only posterior % to %4 finely punctate
while remainder smooth (Fig. 3). In former
Trisoleniella species gaster nearly as long
as high or only very slightly longer, 2nd
abdominal tergite occupies only % of gaster
length and finely punctate (Fig. 8) (shape of
gaster could depend on size of egg mass).
Ventral spine of hypopygium narrow, lon-
669
ger than broad, never reaching beyond apex
of gaster, with very few short scattered pale
setae never reaching beyond apex of spine.
Unisexual females larger than bisexual
ones.
Male. Only known male is that of E. flor-
idana. Antenna 17-segmented (in some
specimens indistinct suture suggests 18th
segment), first three segments flattened, F1l
slightly longer than pedicel and scape to-
gether, strongly curved and enlarged pos-
teriorly. All subsequent flagellomeres, ex-
cept last one, nearly of same length, slightly
longer than broad (Fig. 5). Fore wing mar-
gins with cilia, veins thick and brown. Gas-
ter as long as head and thorax together; 2nd
abdominal tergite occupies entire length of
gaster, subsequent tergites hidden under it.
Otherwise, males similar to females.
Distribution.—Eastern and midwestern
United States. Three species are known
only from Florida: E. floridana, E. enigma,
and E. saltata. Eumayria brevicornis and E.
bignelli are known from New York and
New Jersey; the galls they induce are un-
known.
Biology.—Eumayria floridana is known
only from a bisexual generation and induc-
es stem swelling-like galls, while the four
other species have only unisexual genera-
tions.
KEY TO SPECIES OF EUMAYRIA AND
EXUMAYRIELLA
1. Fully winged; 2nd abdominal tergite with felt-
like ring of dense short pale setae at base, in-
terrupted dorsally
— Brachypterous females; without ring of dense
short pale setae at base of 2nd abdominal ter-
SIS AR eh ewe eee oe ewe te 2
. Frons without or with very few short scattered
white setae; scutum and scutellum slightly
elongated; notauli reaching pronotum; F1 equal
or slightly longer than pedicel and scape to-
gether; small species, length 2.3—3.2 mm. .. .
J hispaps< Sbiynyebawaus oe Bhs Eumayriella invisa (Weld)
— Frons with dense white, long setae; scutum and
scutellum rounded, nearly as broad as long; no-
tauli present at least in posterior half, the sculp-
ture of scutum hidden under dense white pu-
bescence; FI distinctly longer than pedicel and
N
670
scape together; larger species, length 4.0—5.2
100050 eS Gao oneness Eumayriella archboldi, n. sp.
3. Male; Fl curved and broadened in its posterior
Walliffcs ect ng ecec se oee ee Eumayria floridana Ashmead
=iRemale™ oa Sue omen eee ele cee re ene a
4. Head broader than thorax, 1.5—1.7 times broad-
er than long from above; scutum finely aluta-
ceous; antenna 14-segmented; small species
sued itt ameees wea rhage Eumayria floridana Ashmead
— Head as broad as thorax, 2.5—2.8 times broader
than long from above; scutum finely alutaceous
or densely macroreticulate; antenna 14—16-seg-
mented; large specimens, at least 4—5 mm. in
5. Antenna 14-segmented; scutum densely and
uniformly punctate; body black or dark brown
rarite. opie ols Eumayria brevicornis (Beutenmueller)
— Antenna lis—lolsesmented =. es ene 6
6. Antenna 15-segmented; known to induce root
AUS) Fy vayisaees ais: Senne Eumayria enigma (Weld)
— Antenna 16-segmented
8. Scutum very finely and uniformly coriaceous
Sa eas erin arn Eumayria saltata (Ashmead)
— Scutum with much more dull sculpture at basal
end . .Eumayria bignelli (Dalla Torre and Kieffer)
Eumayria bignelli (Dalla Torre and
Kieffer), NEW COMBINATION
Trisolenia punctata Ashmead 1896: 129.
Preoccupied in Andricus by Bignelli
1892. Types examined.
Andricus bignelli Dalla Torre and Kieffer
1902: 61. New name for Trisolenia punc-
tata Ashmead.
Amphibolips montana Beutenmueller
1913a: 122; Weld 1951: 644 (synonym of
punctata). Type examined.
Trisoleniella punctata: Weld 1951: 644
(secondary homonym).
Diagnosis.—Closely resembles E. saltata
in having 16-segmented antenna, however,
basal end of scutum with duller sculpture
than that of E. saltata.
Distribution.—New York, New Jersey.
Biology.—Unisexual females only are
known. The galls and the host associations
are unknown.
Eumayria brevicornis (Beutenmueller),
NEW COMBINATION
Andricus brevicornis Beutenmueller 1913b:
245. Type examined.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Trisoleniella brevicornis: Weld 1951: 644.
Diagnosis.—Closely resembles E. saltata
and E. enigma, however, antenna 14-seg-
mented while in E. enigma and E. saltata
antennae 15—16 segmented; scutum densely
and uniformly punctate; body black or dark
brown.
Distribution.—New Jersey.
Biology.—Unisexual females only are
known. Galls are unknown. Observed ovi-
positing in the buds of Quercus alba L.
(Beutenmueller 1913b).
Eumayria floridana Ashmead
Eumayria floridana Ashmead 1887: 133,
147. Females, males, galls. Types exam-
ined.
Eumayria multiarticulata Ashmead 1887:
133 (name proposed unnecessarily; de-
scription of galls according to those of E.
floridana); Dalla Torre and Kieffer 1910:
601; Weld 1921: 230 (as a synonym of
E. floridana); Burks 1979: 1107 (in un-
placed species of Cynipoidea).
Diagnosis.—Head broader than thorax,
only 1.5—1.7 times broader than long from
above, antenna 14-segmented, while in all
other species of genus head as broad as tho-
rax and 2.5—2.8 times broader than long
from above, and antenna 15—16 segmented.
On basis of galls only, difficult to distin-
guish from some Loxaulus galls and those
of Bassettia floridana. Typically galls of E.
floridana are larger swellings than those
produced by above-mentioned species.
Reared adults and knowledge of host oak
are critical to correct identification.
Distribution.—Indiana, [linois, Arkan-
sas, Texas, Virginia, Florida.
Biology.—Only a bisexual generation is
known. Induces stem-swelling galls at the
base of young sprouts of Quercus coccinea
Muench., Q. falcata Michx., Q. ilicifolia
Wangenh., Q. incana Bartr., Q. laurifolia
Michx., Q. myrtifolia Willd., Q. rubra L.,
Q. texana, Q. velutina Lam. (Weld 1921,
1959). Adults emerge in May through Au-
gust.
VOLUME 99, NUMBER 4
Eumayria enigma (Weld),
NEW COMBINATION
Callirhytis enigma Weld 1921: 219. Fe-
males and galls. Types examined.
Trisoleniella enigma: Weld 1951: 644.
Diagnosis.—Similar to E. saltata, how-
ever female antennae with 15 and not 16
segments. Location of galls and host oaks
different for the two species.
Taxonomic comments.—Morphological
characteristics of the adults suggest that E.
enigma could be a unisexual generation of
E. floridana. It differs from floridana in that
the head as broad as the thorax, while in
floridana the head is broader than the tho-
rax. The only other morphological differ-
ences between these two species are the
number of antennal segments (14 in E. flor-
idana and 15 in E. enigma), and the shape
of the gaster and particularly that of the 2nd
abdominal tergite. Otherwise they are iden-
tical.
Distribution.—INllinois, Texas, Virginia,
Louisiana, Florida.
Biology.—Unisexual females only are
known. Induce root galls on Quercus laevis
Walt., O. myrtifolia, Q. nigra L., Q. rubra,
Q. texana (Weld 1921, 1959). Adult emer-
gence date is unknown. Adults can be dis-
sected from galls by the beginning of No-
vember in Florida (Weld 1921).
Eumayria saltata (Ashmead),
NEw COMBINATION
Andricus (Trisolenia) saltatus Ashmead
1887: 142. Females and galls. Types ex-
amined.
Trisolenia saltata: Ashmead 1903: 155.
Andricus saltatus: Dalla Torre and Kieffer
1910: 549.
Trisoleniella saltata: Weld 1951: 644.
Diagnosis.—The galls are very specifi-
cally shaped, easily distinguished by the
gall alone. For adults, see the key to the
species and diagnosis of E. bignelli, E.
brevicornis.
Distribution.—North Carolina, Florida
(Jacksonville, Ocala - Weld 1926, 1951).
671
Biology.—Unisexual females only are
known. Induces bud galls on Quercus fal-
cata (Weld 1951), Q. laevis, Q. incana
(Ashmead 1887; Weld 1951, 1959), Q. rub-
ra (Weld 1926). Galls in March - April,
adults in April (Ashmead 1887, Weld
1926).
SPECIES TRANSFERRED FROM EUMAYRIA
Andricus longipennis (Ashmead),
NEW COMBINATION
Neuroterus longipennis Ashmead 1887:
140. Type examined.
Eumayria longipennis: Weld 1951: 644.
Comments.—The type for this species is
a male and bears the labels “Jacksonville,
Fla.”’, ‘Collection Ashmead’’, red label
“Type No. 2873 USNM”’, and a handwrit-
ten label ‘“‘Neuroterus longipennis Ashm.”
Ashmead described the female and the gall
“from eight specimens bred May, 1886”
(Ashmead 1886). The description, given by
Ashmead (1887) does not agree with this
specimen. Yet, this is the only specimen of
this species that we were able to find in the
USNM collection. However, L. Weld indi-
cated in his personal catalog (a copy of
which was kindly sent to us by Dr. R. J.
Lyon) that, in addition to the type, there
were two wasps and one gall in an old case
in the USNM and that the collection of
American Entomological Society had 3
wasps. In his catalog L. Weld wrote: *“Type
in USNM has head massive, not broadened
behind eyes, no malar groove, ant. 16-seg.
..’—these characters were the basis on
which Weld moved the species into Eumay-
ria. The term “‘head from above massive’’,
which Weld (1952) defined as a head with
a ‘“‘length at least half width’ from above,
is too imprecise for taxonomic purposes.
For instance, there are bisexual Andricus
species (A. crispator Tschek, A. quercus-
calicis Burgds, A. quercuspetiolicola
(Bass.)) and several North American Cal-
lirhytis species known to have “massive
head’, 1.8—2.0 times only broader than
long. A precise ratio of length and width
672
must be given to avoid confusion. Eumay-
ria floridana has a head only 1.5—1.7 times
broader than long, which can never be
found in Andricus or Callirhytis species.
The two other characters—genae not broad-
ened behind eyes, no malar groove—are
present in many genera. Antenna of the
male of N. longipennis type is 16-segment-
ed, while in E. floridana it is 17- or even
18-segmented. The presence or absence of
the tooth on tarsal claws, as we mentioned
above, is likely a homoplasy; consequently,
it is a character of no generic importance.
An examination of Neuroterus longipennis
type (male) showed that it is not a Eumay-
ria species. It belongs instead to the genus
Andricus in a group of species known to
induce stem-swelling galls. Several North
American species from the current Calli-
rhytis genus eventually must be transferred
to this group as well.
Distribution.—Florida (Jacksonville)
(Ashmead 1887, Weld 1951).
Biology.—Only the male is known. In-
duces stem-swelling galls (Ashmead 1887)
on Quercus laurifolia (Ashmead 1887) and
Q. phellos L. (Weld 1959). Adults emerge
in May.
Eumayriella Melika and Abrahamson,
NEw GENUS
Type species.—Eumayria invisa Weld
(1952b). Holotype (No. 60123) and six
paratypes in the USNM [examined].
Etymology.—The genus is named in hon-
or of Dr. Gustav Mayr.
Diagnosis.—Fore wing never reaching
beyond apex of gaster. Closely resembles
brachypterous species of Trichoteras Ash-
mead: T. coquilletti Ashmead and T. tubi-
faciens (Weld). However, in Eumayriella,
thorax flattened dorso-ventrally; pronotum
dorsally much longer, placed in same plane
as scutum; scutum and scutellum pubes-
cent, each longer than broad, scutoscutellar
suture distinct, scutellum without foveae;
head broader than thorax from above, 2.3—
3.0 times broader than high; antenna fili-
form, F2 shorter than Fl; while in Tricho-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
teras thorax arched in anterior one-third;
pronotum dorsally much shorter and not in
same plane as scutum; each scutum and
scutellum as long as broad; head narrower
or equal to thorax; antennae short, F2 near-
ly equal Fl. Eumayriella also closely re-
sembles Eumayria, but head more trans-
verse in front view and from above (2.3—
3.0 times broader than high, while in bisex-
ual Eumayria only 1.5—1.7 times and in
unisexual Eumayria—2.5—2.8 times; malar
space without radiating striae; antenna fili-
form, long, 14-segmented, F1 substantially
longer than pedicel and scape together; all
flagellomeres much longer than broad. Scu-
tum and scutellum longer than broad, with
dense white setae, without median and an-
terior parallel lines; distinctly emarginated
laterally and partially posteriorly too; scu-
tellum without foveae, with transverse de-
pression along scutoscutellar suture; apical
one-third of scutellum gradually depressed
toward apex and narrowed into point that
joins scutum along median dorsal part;
while posterior one-third of scutellar disk
highest part, strongly convex; sculpture of
scutellar disk very finely punctate, posterior
one-fourth rugose. In Eumayria head nearly
as high as broad, malar space and partially
frons with radiating striae; antennae much
shorter, Fl as long as pedicel, scape as
broad as long; F2 to F4 slightly longer than
broad, subsequent flagellomeres, except last
one, subequal, nearly as broad as long; scu-
tellum with foveae, posterior part of disk
never convex; posteriorly rounded and with
dull rugose sculpture; scutum and scutellum
bare, without dense white setae. Second ab-
dominal segment in Eumayriella without
pale felt-like ring of dense short setae at
base, while Eumayria with such ring. Last
character also used for separation of several
genera of Eucoilidae (Cynipoidea) (Quinlan
1986).
Description.—See descriptions of E. in-
visa and E. archboldi.
Distribution.—Florida.
Biology.—Only females are known.
VOLUME 99, NUMBER 4
Galls in a form of chambers, hidden under
the bark of twigs.
Eumayriella archboldi Melika and
Abrahamson, NEW SPECIES
(Figs. 10-12)
Description. Unisexual female. Entire
body red brown, except for black proepi-
sternum and propodeum. Head from above
much broader than thorax, 3 times broader
than long; in front view transverse (Fig.
10). Occiput, vertex, gena, and frons uni-
formly finely punctate, frons with dense
short pale setae. POL only slightly longer
than OOL; gena broadened behind eye,
nearly as broad as diameter of eye; malar
space uniformly finely punctate, without
groove and radiating striae; nearly 4 times
shorter than length of eye; clypeus de-
pressed, rounded anteriorly. Antenna long,
filiform, 14-segmented in holotype, how-
ever indistinct suture in paratypes suggests
15th segment. Pedicel and scape flattened,
Fl longer than pedicel and scape together,
subsequent flagellomeres distinctly longer
than broad, gradually shortened toward end
(Fig. 11). Pronotum in median dorsal line
longer than usual, only 7 times shorter than
length of scutum; in same range as scutum,
not arched. Sides of propodeum densely pu-
bescent, distinctly visible dorsally, much
broader than usual in Cynipini (Fig. 12).
Scutum smooth, shiny, with areas very fine-
ly coriaceously sculptured, especially lat-
erally, densely pubescent; as broad as long,
with notauli reaching pronotum (in type
specimen) or present at least in posterior
two thirds. Scutum without median and an-
terior parallel lines; emarginated laterally.
Scutellum emarginated posteriorly and lat-
erally, with very indistinct foveae; finely
coriaceous, densely pubescent, distinctly
longer than broad; posterior one-third of
scutellar disk highest part, strongly arched.
Fore wing narrow, barely reaching apex of
gaster, with thick brown veins, with cilia on
margins. Legs slightly lighter than body,
uniformly brown. Propodeum black, medi-
an part shiny, bare, with strong striae, lim-
673
ited by two distinct carinae converging
gradually and slightly inward. Gaster slight-
ly longer than high, base of 2nd abdominal
tergite without ring of pale setae, occupying
nearly half length of gaster; 3rd to 6th ter-
gites visible and finely punctate dorsally
and dorsolaterally. Ventral spine of hypo-
pygium very slightly reaching beyond apex
of gaster, narrow, needle-like, with very
few and short pale setae, never reaching be-
yond apex of spine. Length 3.1—3.6 mm.
Diagnosis.—Frons with dense white long
setae while in E. invisa without/or with
barely visible, very few, short scattered pale
setae. Scutum rounded, nearly as broad as
long; notauli present at least in posterior
half, sculpture of scutum hidden under
dense white pubescence while in E. invisa
scutum slightly elongated, notauli distinctly
reach pronotum. Fl much longer than ped-
icel and scape together, while in E. invisa
F1 equal or only slightly longer than pedi-
cel and scape together and body much
smaller than in E. archboldi.
Types.—Holotype ¢. Paratypes two Q°.
Type locality. Archbold Biological Station,
Lake Placid, Highlands Co., FL, adults
trapped on 4 January 1988 (coll. Mark Dey-
rup). Holotype in the USNM, Washington,
DC, 2 paratype 2 in the private collection
of G. Melika.
Etymology.—The species named in the
honor of Mr. Richard Archbold, founder of
the Archbold Biological Station.
Distribution.—Florida (Archbold Bio-
logical Station, Lake Placid, Highlands
Co.).
Biology.—Only females are known.
Adults emerge in December—January.
Eumayriella invisa (Weld),
NEW COMBINATION
Eumayria invisa Weld 1952b: 335. Females
and galls. Types examined.
Diagnosis.—See diagnosis to E. arch-
boldi. Difficult to distinguish this species on
the basis of galls alone. Galls similar to
those caused by Callirhytis crypta (Ashm.),
674
Bassettia floridana. Adult wasps are nec-
essary for precise identification.
Comments.—We do not provide a de-
scription of E. invisa because it was de-
scribed precisely by Weld (1952a).
Distribution.—Florida (Carrabelle-type
locality-Weld 1952b; Archbold Biological
Station, Lake Placid, Highlands Co.; Jona-
than Dickinson State Park, Martin Co.).
Biology.—Only a unisexual generation is
known. Induces stem galls in the form of
small larval cells hidden under the bark,
usually without external evidence of galling
on Quercus myrtifolia. Adults were dis-
sected from galls in November (Weld
1952b). Adults emerge in January in south-
central Florida (personal observation).
ACKNOWLEDGMENTS
We express our deepest appreciation to J.
Fitzpatrick for his support of our work at
the Archbold Biological Station, M. Deyrup
for providing laboratory space and support,
and C. Abrahamson for extensive field and
technical assistance. We also thank R. Lyon
for suggestions and comments. We thank J.
Abrahamson, R. Bowman, G. Cs6ka, R.
Hammer, A. Johnson, I. Kralick, R. Peet, R.
Roberts, P. Schmaltzer, A. Schotz, C. Wine-
garner, and M. Winegarner for field and
technical assistance. Our special thanks to
A. S. Menke for his valuable suggestions
and support during our work in the National
Museum of Natural History, Smithsonian
Institution (USNM), Washington, DC. Sup-
port was provided to GM and WGA by
Bucknell University’s David Burpee en-
dowment, the Archbold Biological Station,
a National Museum of Natural History,
Smithsonian Institution grant to GM, and
NSF grant BSR-9107150 to WGA.
LITERATURE CITED
Ashmead, W. H. 1886. Synopsis of the North Amer-
ican Sub-families and Genera of Cynipidae.
Transactions of the American Entomological So-
ciety 13: 59-64.
1887. On the Cynipidous galls of Florida,
with descriptions of new species and synopses of
the described species of North America. Transac-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tions of the American Entomological Society 14:
125-158.
1896. Descriptions of new cynipidous galls
and gall wasps in the United States National Mu-
seum. Proceedings of the United States National
Museum 19: 113-136.
. 1903. Classification of the gall-wasps and the
parasitic cynipoids, or the superfamily Cynipoi-
dea. III. Psyche 10: 140-155.
Beutenmueller, W. 1913a. A new species of Amphi-
bolips (Hymenoptera, Cynipidae). Insecutor Insci-
tiae Mestruus 1: 122-123.
1913b. Descriptions of new Cynipidae.
Transactions of the American Entomological So-
ciety 39: 243-248.
Burks, B. D. 1979. Superfamily Cynipoidea, pp.
1045-1107. In Krombein, K. V., P. D. Hurd, Jr.,
D. R. Smith, and B. D. Burks, eds., Catalog of
Hymenoptera in America North of Mexico. Vol-
ume 1. Symphyta and Apocrita. Smithsonian In-
stitution Press, Washington, DC.
Dailey, D. C. and A. S. Menke. 1980. Nomenclatorial
notes on North American Cynipidae (Hymenop-
tera). The Pan-Pacific Entomologist 56: 170-174.
Dailey, D. C., T. Perry, and C. M. Sprenger. 1974.
Biology of three Callirhytis gall wasps from Pa-
cific slope Erythrobalanus oaks. The Pan-Pacific
Entomologist 50: 60—67.
Dalla Torre, K. W. von and J. J. Kieffer. 1902. Genera
Insectorum, Hymenoptera, Cynipidae. Bruxelles,
P. Wytsman. 84 pp.
Dalla Torre, K. W. von and J. J. Kieffer. 1910. Cy-
nipidae. Das Tierreich. Berlin: Friedlander &
Sohn 24, 891 pp.
Eady, R. D. and J. Quinlan. 1963. Handbooks for the
Identification of British insects. London. VIII (Ia).
81 pp.
Fergusson, N. D. M. 1995. The cynipoid families, pp.
247-265. In Hanson, P. E. and I. D. Gauld eds.,
The Hymenoptera of Costa Rica. Oxford, New
York, Tokyo, Oxford University Press.
Gibson, G. A. P. 1985. Some pro- and mesothoracic
structures important for phylogenetic analysis of
Hymenoptera, with a review of terms used for the
structures. Canadian Entomologist 117: 1395—
1443.
Menke, A. 1993. Notauli and parapsidal lines: just
what are they? Sphecos 24: 9-12.
Quinlan, J. 1986. A key to the Afrotropical genera of
Eucoilidae (Hymenoptera), with a revision of cer-
tain genera. Bulletin of the British Museum (Nat-
ural History) 52, 366 pp.
Ritchie, A. J. and T. M. Peters. 1981. The external
morphology of Diplolepis rosae (Hymenoptera:
Cynipidae, Cynipinae). Annals of the Entomolog-
ical Society of America 74: 191-199.
Rohwer, S. A. and M. M. Fagan. 1917. The type-
species of the genera of the Cynipoidea, or the
VOLUME 99, NUMBER 4
gall wasps and parasitic cynipoids. Proceedings of
the United States National Museum 53: 357-380.
Ronquist, EF and G. Nordlander. 1989. Skeletal mor-
phology of an archaic cynipoid, /balia rufipes
(Hymenoptera: Ibaliidae). Entomologica Scandi-
navica. Supplement 33: 1-60.
Weld, L. H. 1921. American gallflies of the family
Cynipidae producing subterranean galls on oak.
Proceedings of the United States National Muse-
um 59: 187-246.
. 1926. Field notes on gall-inhabiting cynipid
wasps with descriptions of new species. Proceed-
ings of the United States National Museum 68: 1—
Sil
. 1951. Superfamily Cynipoidea, pp. 594-654.
675
In Muesebeck, C. FE W., K. V. Krombein, and H.
K. Townes, eds., Hymenoptera of America North
of Mexico. United States Department of Agricul-
ture, Agricultural Monograph No. 2.
. 1952a. Cynipoidea (Hym.) 1905-1950 being
a supplement to the Dalla Torre and Kieffer mono-
graph—the Cynipidae in Das Tierreich, Lieferung
24, 1910. Ann Arbor, Michigan. Privately printed.
351 pp.
1952b. New American cynipid wasps from
galls. Proceedings of the United States National
Museum 102: 315-342.
1959. Cynipid galls of the Eastern United
States. Ann Arbor, Michigan, Privately printed.
124 pp.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 676-680
SEXUAL DIMORPHISM IN MOUTHPARTS OF BLEPHARONEURA LOEW
(DIPTERA: TEPHRITIDAE)
MARTY CONDON, ANN TRUELOVE, AND LYNN MATHEW
(MC), Department of Biology, Cornell College, 600 First Street West, Mount Vernon,
IA 52314, U.S.A.; (AT), Department of Biological Sciences, University of Maryland Bal-
timore County, Catonsville, MD 21228-5398, U.S.A.; (LM), 114 Bagatelle Road, Dix
Hills, NY 11746, U.S.A.
Abstract.—Adults of Blepharoneura manchesteri Condon and Norrbom (Tephritidae),
which rasp and feed upon living plant tissues, are sexually dimorphic in number and
distribution of robust blade-like pseudotracheal ring tips (= blades). Both sexes have
similar numbers of pseudotracheae, but labella of females have more than twice as many
blades as males. Such sexual dimorphism in labellar rasping surface may be associated
with higher nutritional requirements of females. Although sex differences in nutritional
requirements have been reported for other tephritid fruit flies, ours is the first report of
sexual dimorphism in pseudotracheal characters.
Key Words:
Adult fruit flies in the genus Blepharo-
neura Loew (Tephritidae) abrade and break
plant surfaces and feed on released sub-
stances. This behavior, unknown in any oth-
er fly, varies among species of Blepharo-
neura. Some species feed on surfaces of
young leaves, leaving distinctive patterns of
lacy holes, while others feed on the tips of
pedicels (following abscission of flowers),
or on the surface of fruit, leaving distinctive
traces as scars on the fruit surface (Condon
and Whalen 1983; Condon and Norrbom
1994). While feeding, the abdomen of the
fly becomes swollen with fluid the color of
the tissue being rasped. Distinctive morpho-
logical characters are associated with this
behavior (Driscoll and Condon 1994).
Labella of Blepharoneura bear rows of
blade-like pseudotracheal ring tips (=blades)
in a rasp-like arrangement (Driscoll and
Condon 1994). Unlike most flies, Blephar-
oneura have rows of two kinds of labellar
pseudotracheal rings and ring tips: rows of
labella, phytophagy, fruit flies, Cucurbitaceae
open rings with brush-like tips, which form
the channel-like pseudotracheae that deliver
liquid to the mouth as in most flies (Elzinga
and Broce 1986); and rows of highly mod-
ified closed rings with blade-like tips, which
are offset from and parallel to the channel-
like pseudotracheae. Blade-bearing rings are
fused along the length of the blades, sug-
gesting that the modified rings function as
braces for blades that rasp surfaces of plants
(Driscoll and Condon 1994). These distinc-
tive blades are a synapomorphy of the clade
(which includes Blepharoneura and two
poorly known Old World genera, Hexapti-
lona Hering and Baryglossa Bezzi) that may
be the most basal clade of the Tephritidae
(Condon and Norrbom 1994; Han and
McPheron 1994).
As a first step toward understanding the
functional significance of labellar blades and
associated feeding behaviors, we looked for
evidence of sexual dimorphism in mouth-
parts. If blades are used to obtain secondary
VOLUME 99, NUMBER 4
anterior
posterior
Fig. 1. Arrangement of pseudotracheae and blade-
like pseudotracheal ring tips (blades) on the labella of
Blepharoneura sp. (drawing based on female specimen
F3, see Table 1). To emphasize blades, brush-like pseu-
dotracheal ring tips are not shown. Labella are shown
in the open position in which they contact surfaces of
plants. Pseudotracheae are numbered from anterior to
posterior.
compounds used by males in courtship, we
expected males to have proportionately more
blades than females. Alternatively, if nutrients
or other chemicals obtained during adulthood
are used in egg production (Tsitsipis 1989;
Hendrichs et al. 1993), we expected females
to have more blades than males. To test these
hypotheses, we examined male and female
specimens of Blepharoneura manchesteri
Condon and Norrbom, a species that feeds on
surfaces of fruit of Gurania spinulosa Cogn.
(Cucurbitaceae) in northern Venezuela.
MATERIALS AND METHODS
Using specimens of B. manchesteri
reared from seeds of G. spinulosa collected
in Guatopo National Park in northern Ven-
ezuela, we followed slightly modified tech-
niques of Driscoll and Condon (1994) to
prepare labella. We soaked heads of speci-
mens in distilled water for 2—3 days, then
boiled specimens briefly in dilute solutions
677
of KOH or NaOH. After boiling specimens
in distilled water, we allowed the water to
cool to room temperature, and added a few
drops of formaldehyde to the cooling water.
Once mouthparts were everted, we prepared
specimens for SEM using hexamethyldisi-
lanzane (HMDS). Dried specimens were
sputter coated with 16 nm of gold and ex-
amined with a Hitachi 2460-VP scanning
electron microsope.
Beginning with the most anterior pseu-
dotracheae, we counted and numbered
pseudotracheae on each labellar surface,
and counted blades on each pseudotrachea
(Fig. 1). We rotated specimens to get ac-
curate counts because mouthparts were not
always evenly everted.
RESULTS
Labella of male and female specimens
of B. manchesteri differ strikingly in the
number and distribution of blades (Fig. 2,
Tables 1, 2). Despite small sample sizes,
we found statistically significant differ-
ences between males and females in num-
ber of pseudotracheae with blades, num-
ber of blades per pseudotrachea, and total
number of blades on labella (Table 2). Fe-
males had nearly three times as many
blades (92-102) as males (31—44), and
blades were located along twice as many
pseudotracheae. Males and females dif-
fered not only in absolute numbers of
blades, but also in proportion of pseudo-
tracheae associated with blades: blades
were associated with an average of 48%
of pseudotracheae in females, and 23% in
males (Table 2). Males and females did
not differ significantly in total number of
pseudotracheae (Table 2).
In both sexes, blades were concentrated
among the more anterior pseudotracheae
(Fig. 1; Table 1). In males, blades oc-
curred exclusively along anterior pseudo-
tracheae: all posterior pseudotracheae in
males were brush-bordered (i.e. blade-
less). In females, blades occurred along
no more than a third of the posterior pseu-
dotracheae. In both sexes, the posterior
678
Table 1. Distribution of blades along pseudotra-
cheae of individual specimens of Blepharoneura man-
chesteri. Pseudotracheae are numbered from anterior
to posterior (see Fig. 1). A dash indicates absence of
a pseudotrachea. R = right labellum. L = left label-
lum. Specimens are identified by a number preceded
by F (female) or M (male). To highlight anterior to
posterior asymmetry of blade distribution on labella,
bold type indicates presence of blades; a line indicates
the halfway point, half of the pseudotracheae are above
(anterior) to the line, and half are below (posterior) to
the line.
Female Specimens
F3 F9 F38 F45
Pseudo- ———— ee Se
trachea
lm
v2)
I=!
vs)
l=
vs)
It)
v2)
#1
tt
“I
OCOooon Nu 66 GHWOSO
loocooonmmecaac'|
pt peek ek |
—_—- © UO ©
= _
N —)
—_ =
io.)
— mt
| a — ee
—_ |
nNonoo
ot
pS
|
iss ereonmysy ne
ocooooo wm @le
#15 —
Total 47 45
a S30. 54
aS
Rloccoco oOo BNweeecco
Wwoooocooonle 6
Nn
BSS
Nn
Nn
Male Specimens
M10 M19 M35 M37
&
COCOCOCOCOCOIANNOOCSCO
Noocoooooianunnooono
lcococociawzeocnc|
loccoocolawxusccco
cococoooCOoOMmMAaAacCS
coooooaNwoooco
a bocce o clauuwo Se |
loccoocomanocoo
Total 1
N
io)
NO
\O
—
\O
Nn
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Blepharoneura manchesteri, right labella of
male (a, specimen F38) and female (b, specimen M19)
specimens (see Table 1). Labella are swollen beyond
the “‘normal” contact position so that all pseudotra-
cheae are visible. Pseudotracheae without blade-like
pseudotracheal ring tips (= blades) are bordered on
either side by brush-like pseudotracheal ring tips (=
brushes). These brush-bordered pseudotracheae are
densest posteriorly. Mop-like brushes also occur me-
dially (toward the prestomum) on both sides of pseu-
dotracheae that bear blades. Along the portion of the
pseudotrachea where blades occur, brushes are present
only along the posterior margin. Posterior and medial
arrangements of brushes suggest that brushes function
as mops that sop up liquids released by damaged plant
tissue.
portion of the labella usually bears at least
50% more brush-bordered (i.e., blade-
less) pseudotracheae than the anterior por-
tion of the labella (Table 1).
DISCUSSION
We found significant sexual dimorphism
in number and distribution of blades on la-
VOLUME 99, NUMBER 4
679
Table 2. Sexually dimorphic traits of labella of Blepharoneura manchesteri. Summary statistics (Mann Whit-
ney U test) for samples described in Table 1: * = P < .05, ** =P < .025.
Male Female
N =4 N =4
Trait Mean Range Mean Range jz)
Number of pseudotracheae (both labella) 26 25—28 26.3 25—29 NS
Number of pseudotracheae with blades (both labella) 6 6 W2EZ5 11-13 ee
% pseudotracheae with blades 23% 21-24% 48% 41-52% **
Mean number of blades per pseudotrachea with blades 6.1 5.3—7.3 8.3 7.1-9.6 s
Maximum number of blades per pseudotrachea 7.8 7-9 11.25 9-13 S
Total number of blades 36.5 31-44 100.7 92-107 **
bella of B. manchesteri (Fig. 2; Tables 1,2).
Although both male and female flies rasp
plant surfaces, the greater number of blades
in females suggests that they are able to
cause more damage to a greater surface area
of plant tissue than males. The anterior con-
centration of blades and brushes in both
males and females suggest that the mecha-
nism of rasping is similar for males and fe-
males (Table 1). The anterior portions of the
labella probably make contact with the
plant surface first and are moved forward
so that tips of blades contact plant tissue.
In both sexes, the brush-like pseudotracheal
ring tips, which are densest nearest the
mouth and along the more posterior pseu-
dotracheae, probably act as mops to sop up
fluids released by abraded plant tissues
(Fig. 2).
That females have more blades than
males is consistent with the hypothesis that
the unusual plant-rasping behavior of
Blepharoneura is associated with different
nutritional requirements of males and fe-
males. Nutritional requirements of males
and females are known to differ in some
economically important tephritids (Tsitsipis
1989). In some tephritids, ingestion of nu-
trients other than carbohydrates during
adulthood appears to be necessary for pro-
duction of eggs, but not spermatazoa (Hen-
drichs et al. 1991, 1993). Because Blephar-
oneura rasp plants in the Cucurbitaceae,
which often contain cucurbitacins— highly
bitter compounds (Tallamy and Krischik
1989), rasping behavior also could be as-
sociated with aquisition of secondary com-
pounds that could be used as defenses of
larvae or adults.
Are Blepharoneura the only tephritid
fruit flies with sexually dimorphic labellar
surfaces? Despite evidence for sexual dif-
ferences in nutritional requirements in eco-
nomically important tephritids, little is
known about the functional morphology of
their mouthparts. Mediterranean fruit flies
(Ceratitis capitata (Wiedemann)) have
blade-like pseudotracheal ring tips like
those of Blepharoneura, but the blades are
smaller and the blade-bearing pseudotra-
cheal rings are open (like most flies) not
highly modified as in Blepharoneura (Dris-
coll and Condon 1994). In contrast to
Blepharoneura and Ceratitis, Anastrepha,
Bactrocera, and Rhagoletis lack ‘‘blades”’
and have a scalloped arrangement of pseu-
dotracheal ring tips that may function as fil-
ters, not rasps (Elzinga and Broce 1986;
Driscoll and Condon 1994). We could not
find any studies reporting sexual dimor-
phism (or lack of) in labellar surfaces of
other tephritids. Further study of labellar
morphology and feeding mechanisms could
lead to improvements in pest control pro-
grams that target foraging adults, particu-
larly females (R6ssler 1989).
ACKNOWLEDGMENTS
We thank C. Driscoll for his observations
suggesting that Blepharoneura were sexu-
ally dimorphic and for teaching us how to
prepare mouthparts. We thank C. Driscoll,
680
G. Grimes, and P. Rutledge for help with
scanning electron microscopy and photog-
raphy. We thank N. Meyer for graphics, and
G. Steck for helpful comments. Our work
was supported in part by NSF HRD
91-03322, Hofstra University, and the New
York Academy of Sciences.
LITERATURE CITED
Condon, M.A. and A.L. Norrbom. 1994. Three sym-
patric species of Blepharoneura (Diptera: Tephrit-
idae) on a single species of host. (Gurania spi-
nulosa, Cucurbitaceae): new species and new tax-
onomic methods. Systematic Entomology 19:
279-304.
Condon, M.A. and M.D. Whalen. 1983. A plea for
herbivore and pathogen damaged material. Taxon
32: 105-107.
Driscoll, C.A. and M.A. Condon. 1994. Labellar mod-
ifications of Blepharoneura (Diptera: Tephritidae):
neotropical flies that damage and feed on plant
surfaces. Annals of the Entomological Society of
America 87: 448-453.
Elzinga, R.J. & A.B. Broce. 1986. Labellar modifi-
cations of Muscomorpha flies (Diptera). Annals of
the Entomological Society of America 79: 150—
208.
Han, H-Y. and B. McPheron. 1994. Phylogenetic
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
study of selected tephritid fruit flies (Insecta: Dip-
tera: Tephritidae) using partial sequences of the
nuclear 18S ribosomal DNA. Biochemical Sys-
tematics and Ecology 22:447—457.
Hendrichs, J., B.I. Katsoyannos, D.R. Papaj, and R.J.
Prokopy. 1991. Sex differences in movement be-
tween natural feeding and mating sites and trade-
offs between food consumption, mating success
and predator evasion in Mediterranean fruit flies
(Diptera: Tephritidae). Oecologia (Berlin) 86:
223-231.
Hendrichs, J., C.R. Lauzon, S.S. Cooley, and R.J. Pro-
kopy. 1993. Contribution of natural food sources
to adult longevity and fecundity of Rhagoletis po-
monella (Diptera: Tephritidae). Annals of the En-
tomological Society of America 86: 250-264.
Rossler, Y. 1989. Insecticidal bait and cover sprays,
pp. 329-336. In Robinson, A.S. and G. Hooper,
eds. Fruit flies: their biology, natural enemies, and
control. World Crop Pests, Volume 3B. Elsevier,
Amsterdam.
Tallamy, D.W. and V.A. Krischik. 1989. Variation and
function of cucurbitacins in Cucurbita: an exam-
ination of current hypotheses. American Naturalist
133: 766-786.
Tsitsipis, J.A. 1989. Nutrition. Requirements, pp.
103-119. In Robinson A.S. and G. Hooper, eds.,
Fruit flies, their biology, natural enemies, and con-
trol. World Crop Pests, Volume 3A. Elsevier, Am-
sterdam.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 681-692
FOUR PUZZLING NEW SPECIES OF MECOPTERA
GEORGE W. BYERS
Department of Entomology, University of Kansas, Lawrence, KS 66045-2601, U.S.A.
Abstract.—New genus Eremobittacus and new species Eremobittacus spinulatus
(Mexico), Nannobittacus dactyliferus (Eucador), Panorpa sentosa, and Panorpa truncata
(Mexico) are described and illustrated. The unusual characteristics of each species are
discussed.
Key Words:
For several years, the species discussed
here have remained unnamed and unde-
scribed in the hope that additional speci-
mens would be found, of these or closely
related species, that would help to clarify
their relationships with others in their re-
spective genera. This hope, however, has
not been realized. Each of the four species
possesses a striking peculiarity that sets it
uncommonly far apart from its supposedly
nearest relatives. In one case, it was not
possible to assign the species to any exist-
ing genus. The unique and puzzling struc-
tural characters are discussed following the
description of the respective species.
Holotypes, allotypes and most paratypes
are in the Snow Entomological Collection,
Natural History Museum, University of
Kansas, Lawrence, Kansas.
Venational abbreviations, other than the
usual ones of the Comstock-Needham sys-
tem, are as follows: Scv—subcostal cross-
vein from Sc to R, ORs—origin of Rs from
R, FRs—first fork of Rs, Pev—pterostigmal
cross-vein(s), OM—origin of M from Cu,,
FM—first fork of M.
Eremobittacus Byers, NEW GENUS
Similar in many characteristics to Bitta-
cus but differing in (1) length of hind ba-
Mecoptera, Panorpidae, Bittacidae
sitarsus compared to that of fourth tarso-
mere, (2) wing venation, (3) surface sculp-
ture and (4) body colors. In hind tarsus, bas-
al segment approximately same length as
fourth and shorter than second and third to-
gether. Cross-veins between R, and M, in
transverse-diagonal alignment; these outer
three ranks of cross-veins crossing five cells
and conspicuously bordered by clouding
darker than wing membrane generally.
Hairs on most body surfaces very short and
arising from apices of microscopic spinules
(Fig. 7), but spinules much more numerous
than those bearing hairs (especially notice-
able on generally glossy hind femora). Col-
ors contrasting, particularly on legs (Fig. 6);
not seen in other regional Bittacidae.
By existing keys, Eremobittacus will be
identified as the Australian Harpobittacus
because of the relative lengths of the hind
tarsomeres. Transverse-diagonal alignment
of most major cross-veins into three ranks
is not unique (occurs rarely in Bittacus) but
is conspicuous in Eremobittacus due to
dark coloring along these cross-veins. Pres-
ence on much of the body surface of tiny
subconical denticles, or spinules (Fig. 7),
has not been seen elsewhere in the Bitta-
cidae. Mexican species of Bittacus are rath-
er uniformly brown or yellowish brown and
682
do not have the contrasting coloration as in
Eremobittacus.
Type species: Eremobittacus spinulatus,
new species.
Etymology: The generic name is from
the Greek words eremos, solitary or lonely,
and its derivative eremia, desert or wilder-
ness, plus Bittacus. For years known only
from the single specimen described below,
despite efforts to obtain additional represen-
tatives, Eremobittacus is indeed solitary or
alone. The habitat, when I visited it in 1969
and 1972, was semi-desert, along a dry
stream bed bordered by sparse, thorny aca-
cia-like trees and herbaceous plants 1—3 feet
high shaded by these trees; the soil was dry,
stony and mostly bare between the larger
plants. (While the search for additional Er-
emobittacus was unsuccessful on both vis-
its, a still undescribed species of Bittacus
was found, in 1972, in the herbaceous veg-
etation.)
Eremobittacus spinulatus Byers,
NEW SPECIES
(Figs. 1, 6, 7-12)
Description.—Based on 1 d, pinned.
Head: Dorsal surface dark yellowish
brown with extremely short, slender, yel-
lowish hairs at each side above eyes, each
hair arising from a microscopic, subconical
cuticular spinule (cf. Fig. 7); broad median
zone behind ocellar triangle without hairs
(spinules only). Ocellar triangle dark
brownish black with two black setae above
median ocellus; lateral ocelli twice diameter
of median ocellus. Rostrum dark yellowish
brown, genae dark brown; mouthparts am-
ber brown with curved, yellow setae on tips
of maxillae. Antenna dark yellowish brown
with short yellowish hairs; approximately
20 flagellomeres (separations indistinct be-
yond 12 or 13); length about 6 mm.
Thorax: Pronotum with three rounded,
transverse ridges, unevenly dark yellowish
brown, darkest laterally where ridges con-
verge; anterior ridge with low prominence
at each side of wide, shallow, median emar-
gination, each prominence bearing stout,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-5. Wings of new species of Mecoptera. 1,
Left fore wing of Eremobittacus spinulatus, male ho-
lotype. 2, Right fore wing of Nannobittacus dactylif-
erus, male paratype. 3, Left fore wing of Panorpa sen-
tosa, male paratype. 4—5, Right fore wings of Panorpa
truncata, male paratype (4) and female paratype (5).
black setae (on holotype, 2 on left side, 3
on right). Mesonotum and metanotum sor-
did dark yellowish brown with numerous
tiny spinules in broad median band, short
yellowish hairs on each scutellum. Pleural
surfaces, coxae and mera shiny black ex-
cept brownish black on propleuron and
close beneath wing attachments, spinulose,
with sparse, pale setae. Three or four thick,
black setae on outer surface of hind coxa;
two smaller black setae on each epimeron.
Fore femur yellowish brown, not swol-
len, with abundant spinules and interrupted
row of black setae on anterior (4 setae),
VOLUME 99, NUMBER 4
Fig. 6. Eremobittacus spinulatus, male holotype,
left lateral aspect.
dorsal (3—4) and posterior (7) sides; middle
femur resembling fore femur but with 9, 7
and 12 setae in uneven rows. Hind femur
mostly black, dark yellowish brown near
outer end, greatly swollen in basal two-
thirds (Fig. 8), abundantly spinulose (Fig.
7), with some spinules bearing short, dark
hairs; black setae on basal one-third, sparse
(2-4) on anterior (outer) surface, sparse
dorsally, more numerous (14—16) in irreg-
ular row on posterior surface. Tibiae yel-
lowish brown with scattered black setae;
spinules in encircling rows, few bearing
short, pale hairs; tibial spurs long, slender.
Basitarsus of fore leg subequal in length to
second and third tarsomeres together, much
longer than fourth; that of middle leg slight-
ly longer than second and third together.
Hind basitarsus approximately same length
as long, thick fourth tarsomere, shorter than
second and third together.
Wings (Fig. 1) faintly tinged with yel-
lowish, stigma light brown; diffuse light
yellowish brown clouding at ORs, FRs, in
costal and subcostal cells, along all three
ranks of cross-veins, at wing apex and in
basal one-third of wing. Sc ending slightly
beyond level of FRs; Scv opposite FRs.
Cross-veins in radial and medial fields in
approximate transverse-diagonal alignment.
Abdomen of male: Terga 2—6 mahogany-
colored (dark reddish brown), unevenly
darker along posterior and lateral margins,
683
with abundant short hairs, each arising from
slightly raised spinule. Corresponding ster-
na nearly black. Terga 7—8 unevenly black-
ish brown; sterna black. Epiandrial lobes
widely divergent (Fig. 11), only about as
long as basistyles, their dorsal and ventral
margins thickened and approximately par-
allel, apex rounded (Fig. 9), lobes light
brown, with numerous spinules on most of
surface, some bearing short hairs; longer
hairs along lower margin, near apex and on
inner surface; small, black, recurved spines
on inner surface at apex and along upper
margin. Cerci short, narrowing toward tip.
Basistyles light brown with numerous setae
longest and darkest posteriorly below base
of aedeagus and along dorsal margin (Fig.
9). Dististyles conspicuous (Fig. 12), with
dark setae in groups along anterior and pos-
terior margins. Aedeagus (Fig. 10) short,
thick in basal two-thirds, abruptly more
slender toward apex; base flanked by
strongly sclerotized penunci.
Measurements: Body length approxi-
mately 13.8 mm.; length of fore wing 13.3
mm.
Type.—Holotype, 6, and only specimen,
collected near Petlalcingo, Puebla, Mexico,
on 21 August 1963, by E D. Parker and L.
A. Stange. The label indicates three miles
north of Petlalcingo, but only a trail goes
north from the town, into mostly desert
habitat. The actual locality (later confirmed
by Lionel Stange) is at a bridge on High-
way 190, three miles northwest of the junc-
tion of this highway and a side road into
Petlalcingo. The type was presented to the
Snow Entomological Collection, Natural
History Museum, University of Kansas, by
Dr. Frank Parker.
Discussion.—Chance discovery of a spe-
cies (or individual) so different that it jus-
tifies placement in a separate genus is not
without precedent in the Mecoptera. But in
such cases, subsequent searches at or near
the type locality have usually yielded ad-
ditional specimens (e.g., Orobittacus obscu-
rus Villegas and Byers, in central Califor-
nia). In the case of Eremobittacus spinula-
684 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
0.5mm
Figs. 7-12. Eremobittacus spinulatus, male holotype. 7, Detail of spinules on femur; note apical hairs at
upper right and lower left. 8, Left hind leg, left lateral aspect. 9, Terminal abdominal segments, left lateral
aspect. 10, Base of aedeagus, posterior aspect. 11, Epiandrial lobes, dorsal aspect. 12, Right dististyle, dorsal
aspect. Scales: a—figs. 8-11; b—fig. 12.
tus, however, repeated collections made at
the type locality and in the apparently cor-
rect season by myself and others have not
rediscovered this species.
The reasons for placing this unusual bit-
tacid in a genus separate from Bittacus have
been discussed above. The specific name
refers to the microscopic spinules on most
of the body surface, a characteristic unique
to this species.
VOLUME 99, NUMBER 4
685
Figs. 13-14. Nannobittacus dactyliferus, male paratype. 13, Right epiandrial lobe (and part of left), dorsal
aspect. 14, Terminal abdominal segments, right lateral aspect. Scale: both figures.
Nannobittacus dactyliferus Byers,
NEW SPECIES
(Figs. 2, 13-14)
Description.—Based on 2 d, pinned.
Head: Occiput yellowish brown, vertex
and frons above antennal bases black, gran-
ular; ocellar prominence black; lateral ocelli
more than three times diameter of median
ocellus; frons below antennal bases brown
medially, paler next to eyes; clypeus glossy
yellowish brown, labrum darker brown at
sides; maxillary palp brown except apical
segment paler; mandible and maxilla yel-
lowish brown. Eyes large, protuberant, con-
verging slightly below antennal bases. An-
tenna yellowish brown, with approximately
18 flagellomeres (indistinct beyond ninth);
antennal length about 5.1 mm.
Thorax: Pronotum dark brown, with
three transverse, rounded ridges; thick an-
terior ridge slightly upturned with low
prominence at each side bearing one long,
slender, hair-like seta, also shorter setae;
short setae on middle and posterior ridges.
Mesonotum glossy blackish brown with
sparse, fine pale setae; metanotum only
about half as long as mesonotum, blackish
brown, glossy except D-shaped median area
below mesoscutellum. Pleural surfaces,
coxae and mera unevenly brownish gray,
finely pubescent, with sparse yellowish se-
tae longest and most dense on anterior cox-
ae. Femora light grayish brown, darkened
at apex; three or four black setae on outer
surface. Tibiae and tarsi dark yellowish
brown, setae black, tibial spurs nearly black
but with short, yellowish hairs.
Wings (Fig. 2) lightly tinged with yel-
lowish brown, veins brown, with undulat-
ing dark brown markings along costal bor-
der to wing apex, with subapical branch
across outer cross-veins and reaching wing
margin in outer cell M,; five small, brown-
ish transverse clouds in cell M between OM
and FM. In fore wing, Scv opposite ORs,
Sc joins C opposite FRs; one Pcv; 1A end-
ing slightly beyond level of h; narrow,
brownish borders around nygmata.
Abdomen of male: Terga 2—7 sordid dark
yellowish brown anteriorly, dark brown
posteriorly; corresponding sterna narrow,
sordid yellowish brown except sternum 7
lighter yellowish brown and abruptly wid-
ened posteriorly. Tergum 8 brown through-
out, with low, rounded dorsolateral lobe at
each side; sternum 8 light brown. Epiandri-
al appendages (Fig. 14) brown along dorsal
and apical margins, including slender, ter-
686
minal (posterior) prolongation which
curves slightly mesad, pale brown ventrally
and pale yellowish brown on slender ven-
tral prolongation; slight protuberance on
dorsal margin near mid-length, directed
mesad, another, rounded and flattened, near
base of ventral prolongation. Black spines
on dorsal protuberance and on inner dorsal
margin of epiandrial appendages (Fig. 13).
Basistyles only about half as long as epian-
drial appendages, brown dorsally grading
into light brown ventrally and posteriorly.
Dististyles inconspicuous, short, thick,
rounded, strongly turned inward and for-
ward. Cerci nearly as long as dorsal edge
of basistyle, sharply pointed at apex, brown
except pale near attachment to proctiger.
Aedeagus long, coiled, unmodified near
base, becoming filiform at approximately
level of lower edge of epiandrial append-
age.
Measurements: Body length 17.0—18.8
mm. (holotype 17.0 mm); length of fore
wing 17.2—18.0 mm. (holotype 17.2 mm.).
Types.—Holotype, 6, collected in Mal-
aise trap, in Sucumbios, Ecuador (0.5°S,
76.5°W), elev. 270 m., 12-22 February
1995, by Peter Hibbs; specimen received by
way of Dr. J. S. Ashe. Paratype d, in Mal-
aise trap, Limoncocha, east of Coca, Napo
Province, Ecuador, 22 May 1976, by David
G. Young; received from Dr. C. P. Alexan-
der, who had found it among Ecuadorian
crane flies sent to him. Habitat for both
these specimens is described as wet, low-
land tropical forest (secondary forest in the
case of the paratype).
Discussion.—In most characters, Nan-
nobittacus dactyliferus is not strikingly dif-
ferent from other known species in its ge-
nus, but the epiandrial appendages (tergum
9) are conspicuously different from those in
any other species. Nannobittacus pollex
Byers and Roggero has a small, thumb-like
projection from the lower margin of each
epiandrial appendage, but this projection is
less than one-fourth the length of those in
dactyliferus. The name dactyliferus is de-
rived from the long, slender prolongations
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
of the epiandrial appendages (Latin dacty-
lus, from Greek dactylos = finger; fero =
to bear, carry). The possible function of the
long, finger-like prolongations, the one at
about a right angle to the other, is problem-
atical.
Panorpa sentosa Byers, NEW SPECIES
(Figs. 3, 15-19)
Description.—Based on 22 ¢ and 27 2,
pinned, and 1 6, 2 @ in alcohol.
Head: Dorsum mostly shiny dark yel-
lowish brown, slightly darker (or usually a
brown spot) at each side of occiput adjacent
to eye; ocellar prominence black, lateral
ocelli about 1.5 times width of median ocel-
lus. Rostrum yellowish brown, maxillary
palps darker, dark brown on apical half of
terminal segment. Antennal scape yellowish
brown, pedicel brown, flagellum brownish
black to black, with 32—36 flagellomeres in
male (holotype 32 and 33), 33—36 in female
(allotype 36). Antennal length, male, about
9.0 mm., female about 7.8—8.0 mm.
Thorax: Pronotum shiny dark brown ex-
cept yellowish brown along mid-line and
widening toward rear; 5—6 black setae
along anterior margin on each side. Meso-
notum and metanotum with broad, pale yel-
lowish brown median stripe, brown at sides,
nearly black just anterior to bases of fore
wings; setae numerous, short, dark. Pleural
surfaces, coxae and mera unevenly yellow-
ish brown with scattered setae longest and
most numerous on anterior surfaces of cox-
ae. Femora, tibiae and tarsi yellowish
brown, tibial setae black, fifth tarsomere
dark brown.
Wings (Fig. 3) lightly tinged with yel-
lowish brown; spots light brown, no com-
plete bands; spots at ORs, FRs, from prox-
imal end of stigma to fork of R,,; or to M,,
outer end of stigma across fork of R,,3; to
R,, in cell Ist R,, second cell M;, cell M,
and second cell Cu,, in outermost radial and
medial cells at wing margin, and along out-
er cross-veins, but variable.
Abdomen of male: Terga 2-5 light yel-
lowish brown with short, pale setae; sterna
VOLUME 99, NUMBER 4
Figs. 15-19.
dististyle, male, mesal aspect. 17, Ninth abdominal tergum, male, dorsal aspect. 18, Aedeagus and parameres,
male, left lateral aspect. 19, Genital plates of female, ventral aspect (posterior end at top). Scales: a—figs. 15—
17; b—figs. 18-19.
2-5 slightly paler. Segments 6—9 yellowish
brown. Notal organ comprising broadly
rounded, mid-caudal margin of tergum 3
with short, downcurved yellow setae, and
sharp, strongly sclerotized peg on anterior
tergum 4. Posterodorsal surface of segment
6 glabrous and slightly depressed (genital
bulb ‘‘closes’”’ against this area). Segments
7 and 8 short, 1.5—2 times as long as their
diameter. Tergum 9 (Fig. 17) expanded at
apex, lateral corners acute, caudal margin
slightly rounded, nearly truncate; cerci dis-
tinctly two-segmented. Sternum 9 (Fig. 15)
prolonged to nearly half length of basi-
styles, then separated into slender hypov-
alves that extend slightly beyond basistyles.
Outer margin of each dististyle slightly con-
cave, apex moderately curved, strongly
sclerotized; basal cup greatly prolonged
ventrad (Fig. 16), its blunt apical margin
687
0.5mm
Panorpa sentosa, paratypes. 15, Genital bulb of male, ventral (posterior) aspect. 16, Left
with single small tooth in most individuals;
blackened, acute spine near base. Aedeagus
(Fig. 18) with undivided ventral and dorsal
parameres, dorsal ones compressed, some-
what spatulate, rounded at apex. Ventral
parameres well sclerotized, rod-like through
most of their length, extending beyond ends
of basistyles and projecting ventrad be-
tween hypovalves, each deflected slightly
laterad near base; apex variable, generally
of two short, flat, expanded arms with spi-
nose margins, often with a few setae. Ven-
tral and dorsal valves small, concealed be-
tween bases of parameres in lateral aspect.
Abdomen of female: Terga shiny yellow-
ish brown to light brown; sterna 2—5 slight-
ly paler than corresponding terga; setae
pale; cerci black. Subgenital plate broadly
rounded posteriorly, with small median
point in some individuals, slightly rounded
688
at sides, keeled along ventral midline. Pos-
terior margin of apical genital plate (Fig.
19) nearly transverse, lateral edges curved
ventrad; basal plate indistinct, only weakly
sclerotized; axial portion greatly elongated,
densely sclerotized through most of its
length, anterior apodemes short, pale.
Measurements: Body length, male, about
7.1-9.2 mm. (holotype 8.8 mm.); female
about 7.3-9.2 mm. (allotype 9.2 mm.).
Length of fore wing, male, 8.7—11.1 mm.
(holotype 10.4 mm.); female, 10.3—11.2
mm. (allotype 10.5 mm.).
Types.—Holotype, ¢, allotype, two ¢
and three 2 paratypes collected 14.9 mi.
(24 km.) west of El Naranjo, San Luis Po-
tosi, Mexico, on 26 August 1972, by G. W.
Byers (field catalogue San Luis Potosi No.
15) and A. R. Thornhill. Additional para-
types, from San Luis Potosi: 14 mi. west of
El] Naranjo, 21 June 1971, N. D. Penny (3
3,5 2); 15 mi. west of El Naranjo, 5 July
1971, (N.'D> Pentiy (7.6; 6 +2); iwy. 70;
km. 82, along road to microwave tower Mi-
croondas Tortugas, 20 July 1988, C. L.
Smith (3 6, 4 2); 16 mi. west of El Na-
ranjo, 3500 ft., 8 September 1992, Wes Bi-
cha (4 6, 1. 2) and 9 Sept. 1992 2 6,7
2); from Tamaulipas, Rancho del Cielo,
3800 ft., 8 mi. west of Gomez Farias, 24—
29 July 1971, G. E. and K. E. Ball (3 d, 4
?). Specimens collected by C. L. Smith are
from the collection of the University of
Georgia, Athens; those collected by Wes
Bicha are in his collection.
Discussion.—Habitat at the type locality
was forest of various oaks and a few other
kinds of trees, beside Highway 80, 14.9
miles by road west of El Naranjo (1.6 miles
below summit of pass and about 31 miles,
or 49.5 km., by road west of Antiguo Mo-
relos). Branches of all large trees bore nu-
merous epiphytic bromeliads, mosses and
liverworts; the undergrowth included
woody shrubs up to two meters in height
and herbaceous plants a meter or less high.
Elevation 4000 feet (1220 m.); temperature
68°F; weather 100% cloudy, with rain end-
ing collecting at 10:40 a.m. This is one of
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
a very few localities in Mexico where Pa-
norpa has been found lower than 5000 feet
(1524 m.), perhaps because this is one of
the northernmost places in Mexico where
the genus has been found.
Norman D. Penny discovered the species,
and information about its occurrence was
passed to Byers and from him to Wes Bi-
cha. Cecil Smith’s finding it was an inde-
pendent event.
The shape of sternum 9 and its hypo-
valves and the projecting ventral parameres
with spinose apices make males of Panorpa
sentosa readily recognizable. It is the ter-
minal structure of the ventral parameres that
gives the species its name (Latin sentosa =
thorny). The unique tergum 9 is often not
easily seen in pinned specimens; three other
Mexican species of Panorpa, in the invo-
luta group (Byers 1996), have the ninth ter-
gum with approximately transverse poste-
rior margin. In size and wing maculation P.
sentosa somewhat resembles P. mucronata
Byers, known only from Hidalgo, but the
characters mentioned will easily differenti-
ate males of these species. Females also can
be readily recognized by the shape of the
subgenital plate, which is short, broad, ven-
trally keeled and with a broadly rounded
posterior margin in sentosa but unusually
long and narrowly rounded posteriorly in
mucronata.
Panorpa sentosa is a puzzling species
because of the male’s long ventral para-
meres that project conspicuously from with-
in the genital bulb and have a complicated
apex with numerous thorn-like points on
the margin. In males of all other Mexican
species, the ventral parameres have a sim-
ple apical margin that may be pointed or
blunt but not irregularly jagged. Also, ex-
cept in P. sentosa, the ventral parameres are
shorter than the dorsal parameres, or in
those species with two-branched ventral
parameres the ventral branch is conspicu-
ously shorter than either the dorsal branch
or the dorsal parameres. When the ventral
parameres project from the genital bulb,
only their apices can be seen in lateral as-
689
VOLUME 99, NUMBER 4
tips
Rat tses
pees _
Genital bulb,
Ninth abdominal tergum, dorsal aspect. 21,
showing truncated ninth sternum. Scale: both figures.
20,
male, paratype.
’
cata
Panorpa trun
ile
ventral (posterior) aspect
Figs. 20
>
black with
sparse, very short, pale hairs on much of
surface
hiny
inence s
lar prom
ing oce
as contrasted to half 1
°
ing ventrad
ies
ther spec
their length proj
Sd.
pect, in o
sento-
in P
ect
clypeus shiny dark yellowish
’
’
maxillary
ibles and max-
llae brown. Scape and pedicel dark brown
brown, labrum unevenly brown
palps yellowish brown, mand
i
39 ¢ flagellum mahogany brown basally, grad
N
a)
=
O
es
a
N
oe
aa
Bo
eal
Pama
<a
2 50
re
3
=
S
S
AY
le
(holotype 44), 41-44 in female. Antennal
ing
in ma
42-45 flagellomeres
°
black
into
-}
3
d, and 3 6,4 @ preserved in alcohol.
Description.—Based on 36
pinne
vertex and frons includ-
b
t
ipu
: Occ
Head
690
length about 11-13 mm. in male, 11—12
mm. in female.
Thorax: Pronotum mostly black, dull yel-
lowish brown medially on posterior trans-
verse ridge, with very short, pale hairs; an-
terior margin turned upward, with 6—8 black
setae and several shorter black hairs at each
side. Mesonotum black at sides, light brown
medially including scutellum; metanotum
black at sides, more broadly light brown me-
dially than mesonotum. Pleural surfaces,
coxae and mera black with fine, short, whit-
ish pubescence and sparse yellowish setae
most dense on coxae and lower parts of
mera. Fore and middle legs orange-brown on
femora and tibiae, tarsi darker, fifth tarso-
mere black. Femur of hind leg dark orange-
brown, tibia sordid yellowish brown, tarsus
brown, darkening toward apex.
Wings tinged with yellowish brown, un-
marked except for slightly darkened stigma,
in male (Fig. 4), with dark brown apical
band, from near end of R, to end of M, and
to wing apex, and slightly darkened stigma,
in female (Fig. 5). Veins R, and R; un-
branched. Whitish thyridium at FM.
Abdomen of male: Segments 2-8 ferru-
gineous with short, pale yellowish hairs
sparse on terga, more dense on sterna; hairs
longer on sixth segment. Segments 7 and 8
each about as long as 6, much longer than
more anterior segments. Notal organ a
broad, truncate median lobe on hind margin
of tergum 3, with downcurved yellowish
hairs, and a low, blunt median process on
tergum 4 with yellowish hairs directed
cephalad. Segment 9 shiny black to brown-
ish black except ferrugineous on petiole in-
serted into segment 8; setae brown to black.
Ninth tergum (Fig. 20) elongate, narrowing
posteriorly to acute apex, with long, black
hairs along sides beyond slender, apparently
single-segmented cerci; ferrugineous spot
near apex. No hypovalves or other prolon-
gations on ninth sternum (Fig. 21). Basi-
styles fused for more than half their length.
Dististyles (Fig. 21) black basally, dark or-
ange-brown in apical half; basal cup usually
with blackened tooth on inner, dorsal mar-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
gin. Aedeagus (Figs. 22, 23) with two-
branched vental parameres; ventral branch
only weakly sclerotized, covered laterally
with short hairs, its ventral prolongation
curved slightly mesad; dorsal branch rod-
like, darkly sclerotized at base, pale at apex.
Dorsal parameres compressed, wide in lat-
eral aspect, rounded at apex. Ventral valves
acutely tipped, more darkly sclerotized
along dorsal (upper) curvature than on low-
er side. Dorsal valves small, densely scler-
otized, apically rounded.
Abdomen of female: Terga 2—5 unevenly
dark ferrugineous to dark brown (probably
some post-mortem effects involved), with
short, pale setae; terga 6—8 orange-brown,
9 dark brown, 10 brown on posterior half,
with long, black setae; cerci black. Sterna
2-5 yellowish orange to light ferrugineous,
wide, with pale setae; 6—8 ferrugineous.
Conspicuous laterotergites, attenuate ante-
riorly, rounded posteriorly, on segments 7
and 8. Subgenital plate (Fig. 24) broad with
transverse or very broadly rounded caudal
margin and long bordering hairs; mostly
dark brown but abruptly paler at base. Gen-
ital plates (Fig. 25) with divergent anterior
apodemes; apical plate white, only weakly
sclerotized; extent of basal plate not evi-
dent.
Measurements: Body length, male, about
20.6—22.4 mm. (holotype 22.4 mm.); fe-
male, about 14.7—15.4 mm. (allotype 14.9
mm.). Length of fore wing, male, 16.6—17.4
mm. (holotype 17.4 mm.); female, 16.3—
17.1 mm. (allotype 16.3 mm.).
Types.—Holotype, ¢, allotype and 19 d,
23 ¢ paratypes collected near Highway
110, 5.4 km. by road north of Mazamitla
(measured from major road junction at
north edge of the town), Jalisco, Mexico,
10 July 1985, at elevation 6890 feet (2120
m.), by George W. Byers (field catalogue
Jalisco no. 5). One ¢, two 2 paratypes
from nearly same locality (a few hundred
meters farther north), collected by David K.
Faulkner, 12 July 1982, in the Natural His-
tory Museum, San Diego, California; 18 ¢
and 17 2 paratypes collected 3.6 mi. (5.8
VOLUME 99, NUMBER 4
0.5mm
Figs. 22-25.
Panorpa truncata, paratypes. 22, Aedeagus and parameres, male, left lateral aspect. 23, Ae-
deagus, ventral aspect. 24, Subgenital plate of female, ventral aspect (posterior end at top). 25, Genital plates
of female, ventral aspect (posterior end at top). Scales: a—figs. 22—23, 25; b—fig. 24.
km.) south of Mazamitla, 26 and 29 August
1989, by Wes Bicha, in his collection.
The type locality, about 30 km. south of
Laguna de Chapala, is a small valley in
sparse oak woods (trees 10 to 13 m. high),
with shrubs 1—2 m. high, lower herbaceous
plants and grasses; slope of valley floor
about 15—20 degrees.
Discussion.—This species was discov-
ered by Dr. David K. Faulkner of the Nat-
ural History Museum, San Diego. In 1984,
he sent me one male and two females for
identification. The first thing one notices
about Panorpa truncata, apart from its
large size, contrasting coloration (Fig. 26),
and the sexual difference in wing colora-
tion, is that there is no prolongation of the
male’s ninth sternum, which in other spe-
cies is usually divided into separate hypo-
valves. Males of every other species of Pan-
orpidae in the world possess this structure.
It has proved to be useful in taxonomy be-
cause it varies in shape from species to spe-
cies but is fairly constant within a species.
Describing Faulkner’s obviously new spe-
cies based upon what appeared to be a mon-
strosity seemed unwise. Accordingly, I
made a brief trip to Jalisco the next summer
(1985) to find additional males. And all of
them lacked hypovalves. The species takes
its name from the absence of these sternal
prolongations (Latin truncata = cut off, or
deformed, mutilated). I have never deter-
mined the function of hypovalves in male
panorpids but assume they are tactile; clear-
ly P. truncata has no need of them.
Fig. 26.
Males of P. truncata, when inactive, hold
their wings roof-like above the body and
the tip of the abdomen curved forward so
as to be virtually concealed by the wings.
Females, more active than males, hold their
black-tipped wings more out to the sides.
Both males and females were often (usu-
ally, if undisturbed) seen resting in a ver-
tical position, such as clutching a plant stem
or tall grass blade. When alarmed, females
flew higher than males into low shrubs and
herbaceous vegetation.
At the time of these observations, two
males of a second (still unnamed) species,
somewhat smaller than P. truncata and with
orange body including orange genital bulb,
were collected. These had well-developed
hypovalves.
ACKNOWLEDGMENTS
I am indebted to David Faulkner, Norman
Penny, Frank Parker, and Lionel Stange for
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Male Panorpa truncata, left lateral aspect. Drawing by Anne Musser.
discovering three of the species described
here, also to Wes Bicha for his tireless ef-
forts to make the Mecoptera better known.
My thanks to all these colleagues and to
Steve Ashe and Cecil Smith for their aware-
ness of my interest in Mecoptera and for
forwarding specimens. The efforts and pa-
tience of Sharon Lee Hopkins and Cynthia
Woods in putting the paper into computer
and making corrections are much appreci-
ated. Comments by two anonymous review-
ers are similarly appreciated. This is con-
tribution number 3188 from the Division of
Entomology (Snow Entomological Collec-
tion), University of Kansas Natural History
Museum, Lawrence, Kansas 66045, U.S.A.
LITERATURE CITED
Byers, G. W. 1996. Descriptions and distributional rec-
ords of American Mecoptera. IV. The University
of Kansas Science Bulletin 55(14): 519-547, 99
figs.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 693-696
BIOLOGICAL NOTES ON SPARASION LATREILLE (HYMENOPTERA:
SCELIONIDAE), AN EGG PARASITOID OF ATLANTICUS GIBBOSUS
SCUDDER (ORTHOPTERA: TETTIGONIIDAE)
E. E. GRISSELL
Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart-
ment of Agriculture, c/o National Museum of Natural History, MRC 168, Washington,
DC 20560 U.S.A.
Abstract.—The first behavioral observations for any species of Sparasion and the first
report of the genus Aflanticus (Orthoptera: Tettigoniidae) as a host of Sparasion are pre-
sented. In Florida, a number of female wasps were observed burrowing headfirst into
sandy areas. In every instance where a female burrowed into sand and the area subse-
quently was excavated, an egg of Atlanticus, oriented vertically, was found at 12 to 15
mm beneath the surface. Females emerged headfirst from the sand if they remained un-
derground for more than a few minutes. A single female was excavated from the ground
while in the process of ovipositing into an egg; she was at its uppermost end with her
head oriented toward the surface.
Key Words:
egg, parasitoid
The genus Sparasion Latreille is repre-
sented by over 100 species throughout the
Holarctic and Oriental regions (Johnson
1992), eight of which occur in America
north of Mexico (Muesebeck 1979). Essen-
tially nothing is known about the biology
or behavior of these wasps. Kozlov and Ko-
nonova (1990) recently described over 50
new species of Sparasion, and not a single
one had been reared. The paucity of bio-
logical or behavioral data for species in this
genus is doubly remarkable because it has
been recognized for nearly 200 years (de-
scribed in 1802), and its single known host
in the Americas is the Mormon cricket, An-
abrus simplex Haldeman, an insect of leg-
endary stature in the United States. Sur-
prisingly, even for this common, well-
known host, no truly biological or behav-
ioral observations have been published for
its parasite, Sparasion pilosum Ashmead,
Hymenoptera, Scelionidae, Sparasion, Tettigoniidae, Atlanticus gibbosus,
except the host record itself (Cowan 1929)
and subsequent citations of this record
(Mills 1941, Hitchcock 1942, Wakeland
1959, Muesebeck 1979). Although a few
additional papers refer to Sparasion in re-
lation to a potential host, these were merely
specimens of Sparasion collected in a hab-
itat relative to the potential host. For ex-
ample, Spencer (1958) reported Sparasion
sp. “probably parasitic upon eggs of the tet-
tigoniid Anabrus longipes Caudell,”’ and
Thorens (1991) collected Sparasion sp. in
a locality containing an acridid grasshopper
(Chorthippus sp.).
In this paper I present anecdotal obser-
vations made on the behavior of an unde-
scribed species of Sparasion in Florida that
attacks eggs of Atlanticus gibbosus Scudder
(Orthoptera: Tettigoniidae). This is the first
report of Atlanticus as a host for Sparasion.
These observations were made in 1974 and
694
1975 and were to form the basis of more
detailed studies to follow. Other obligations
prevented further investigation, however,
and as no one has published biological in-
formation for the genus in the intervening
twenty years, my notes provide enough data
to highlight significant aspects of the here-
tofore unknown behavior of a species of
Sparasion.
METHODS
I made observations in two areas about
16 km apart in Alachua County, Florida. In
both areas the soil consisted of sand, which
in the absence of rain, was loose and pow-
dery dry for the top 1 or 2 cm.
Alachua (0.5 km southeast). This site
was a large, sandy, disturbed area formed
by the intersection of a railroad track and
several dirt roads. The entire area had been
bulldozed through a low hillock in recent
times, probably as a barrow pit. A number
of annual and perennial plants, most nota-
bly Cassia sp. (Fabaceae), were returning
to the site.
Gainesville (grounds of the Florida State
Collection of Arthropods, Division of Plant
Industry building). This site was an un-
paved road that had been cut through the
woods around the back side of the DPI
building. The site has been paved in the in-
tervening years since observations were
first made.
Determination of the status of this Spa-
rasion as undescribed was made first by the
late C. EK W. Muesebeck and subsequently
was confirmed by Lubomir Masner, Agri-
culture Canada. Additionally, I have com-
pared the species with types and other spec-
imens in the National Museum of Natural
History, Smithsonian Institution, and it ap-
pears to differ from all available material.
Currently no systematist in authority is
willing to describe the taxon. All voucher
specimens collected in this study are
housed in the Florida State Collection of
Arthropods, Gainesville, Florida, along
with large series of material representing
the same taxon.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Field recognition of this species requires
experience based upon preserved speci-
mens. The wasps are ca. 5 mm in length,
appear black in color, and are most easily
seen as they fly or walk back and forth over
a small area of soil. Females are distin-
guished from males by the orange legs
(black in males) and short, hidden antennae
(elongate and easily seen in males).
Determination of Atlanticus gibbosus
was made by D. A. Nickle, who compared
dried eggs excavated from the Alachua site
(21 July 1975) with eggs dissected from
dried adult female Adlanticus in the collec-
tion of the National Museum of Natural
History. Atlanticus gibbosus is the only
common shield-backed grasshopper found
in Florida and is particularly abundant dur-
ing the summer months when my obser-
vations were made.
OBSERVATIONS
Alachua, 21 to 27 July 1974.—On 21
July I arrived at the site at 10:45 am and
saw several female Sparasion flying slowly
over a small sandy area at about 2 to 5 cm
above the surface. Two to 5 wasps were
seen walking or flying in any given area
(approximately 30 cm? to 1 m’). Once a fe-
male landed on the ground she walked
about erratically over the surface with her
abdomen bobbing slightly up and down,
wings folded and held horizontally over the
abdomen. The antennae were extended
downward in an inverted V-shaped pattern
just above the surface. When a female
found an area of interest, she touched her
antennae on the sand and vibrated them;
then she would plunge headfirst into the
sand. As she entered, she rotated her body
from side to side, and her antennae ap-
peared to play some role in excavation, but
this could not be confirmed. In most cases
her body would quickly disappear beneath
the surface, and just as quickly she would
back out completely and begin the probing
activity in nearby areas. Females did not
come out headfirst unless they had been un-
derground at least several minutes.
VOLUME 99, NUMBER 4
In one instance I saw a female emerging
headfirst from the sand. I excavated at the
point of emergence and found a tettigoniid
egg ca. 6 mm in length at 13 mm below
the surface (top of egg at 13 mm). A few
minutes later I saw a female wasp burrow
headfirst into the sand. It took 40 to 50 sec-
onds for her to completely disappear from
sight. Five minutes later she emerged head-
first from the soil. Then almost immediately
another female burrowed headfirst into the
sand in the same area. After 8 minutes I
excavated the soil and found the female
near an egg. The top of this egg was 10
mm beneath the surface. The egg was 6 mm
in length. Shortly thereafter I saw another
female burrow headfirst into the ground and
completely disappear in ca. 45 seconds. Af-
ter waiting 17 minutes for her to emerge, I
excavated the sand and found an egg. The
female wasp was oriented head upwards,
above the tip of the egg, with her ovipositor
embedded into it. When I removed the egg,
the wasp was still attached to it, but she
broke free and flew away. This egg was 12
mm below the surface and was 6 mm long.
I placed the above eggs in gelatin capsules,
but nothing emerged from them.
On 23 July I visited the same site from
9:30 to 11:30 am and saw numerous males
flying above the sand; some alighted on the
ground, others alighted on Cassia. I did not
see any females until 10:15 and observed
no digging or mating attempts.
On 25 July I visited the site at 10:30 am.
The sand was slightly damp on the surface.
I saw males and females cruising the area.
In one case I saw 7 females on the ground
within a 30 cm? area. Some began to bur-
row into the sand but then stopped. No fe-
males entered the ground. Some of the fe-
males approached each other, and occasion-
ally one would “‘hop”’ at the other (a short
jump of 1 to 2 cm). Several times I saw
females digging within 5 mm of each other.
It began to rain heavily, and the wasps dis-
appeared.
On 27 July I arrived at 11:00 am. A few
males were flying over the sand and landing
695
on Cassia. The area was extremely wet, and
no females were seen. It rained heavily for
the next few days as well, and I abandoned
the site.
Gainesville, 2 July 1975.—At 10:45 am,
along the edge of a sandy road, I observed
a female burrow headfirst into nearly level,
loose, dry sand. In 10 minutes a female
(presumed to be the same, but perhaps in-
correctly, see below) emerged headfirst
from the sand about 3 mm from where she
entered. Her body was covered with dust,
and she spent several minutes cleaning her
abdomen and then her head. Then she
walked away. I immediately excavated the
sand beginning about 10 cm away from the
emergence site. I excavated to a depth of
20 mm (the first 15 mm were dry, then be-
came damp). At 15 mm in depth and near
where the female emerged, I found two
eggs of a tettigoniid a few cm apart. These
were about 5 mm in length and placed ver-
tically in the moist sand with the tops at the
15 mm level. I also found three female
Sparasion, heads upright, near these two
eggs but not in contact with them. These
females either walked or flew away quickly
as the sand fell away from their bodies dur-
ing my excavation. I brought the eggs into
the lab and placed them in gelatin capsules,
but nothing emerged.
DISCUSSION
During eight days of observations (over
a two year period), the number of both male
and female wasps seen flying and landing
on the sand varied, with up to seven fe-
males in a 30 cm? area. The absence of ob-
served matings was unusual for such an
abundance of individuals, but matings may
have taken place earlier or later than the
period during which my observations were
made (in late morning, ca. 9:30 to 11:30
am).
Females seemed most attracted to areas
of dry, friable sand; wet sand appeared to
pose a deterrent to initial surface penetra-
tion. All observed females entered the sand
headfirst. After a female entered the sand
696
her presence was undetectable from the sur-
face until she either backed out (usually
only a few seconds after penetration) or
emerged headfirst. It appeared that once a
female found a spot beneath the surface that
was of interest, presumably because a tet-
tigoniid egg was present, she managed to
reverse direction under the sand. Females
apparently could detect host eggs from the
surface, but they either could not detect oth-
er wasp females or these females were not
a deterrent.
Although I could not demonstrate by ev-
idence of successful rearing that Atlanticus
gibbosus is, in fact, a true host of this Spa-
rasion, I believe that the observations point
to this conclusion. The constant association
of female wasps with Aftlanticus eggs and
the presence of a female wasp ovipositing
into an egg both suggest that this tettigoniid
is likely to be a host. Additionally, Atlan-
ticus is a member of the Tettigoniinae, as is
the only proven host of Sparasion in the
Nearctic, the Mormon cricket.
ACKNOWLEDGMENTS
I thank David Nickle, Systematic Ento-
mology Laboratory, USDA, Washington,
DC, for identification of the tettigoniid eggs
and for information on tettigoniine tettigo-
niids. For reading and commenting upon
the manuscript I thank Lubomir Masner,
Agriculture Canada (Ottawa), Andy Austin,
The University of Adelaide (Adelaide, Aus-
tralia), John Brown and David Smith (Sys-
tematic Entomology Laboratory, USDA,
Washington, DC), and an anonymous re-
viewer. I am indebted to C. E W. Muese-
beck (deceased) and L. Masner for infor-
mation and help with the identification of
this species. I thank Jim Wylie, Florida
State Collection of Arthropods, for the loan
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
of voucher specimens of Sparasion, which
are housed in the FSCA collection. Finally,
I thank Richard Dysart, European Biologi-
cal Control Laboratory, USDA, for access
to his bibliography of scelionid references,
and Ray Gagné, Systematic Entomology
Laboratory, USDA, for the translation of
some French text.
LITERATURE CITED
Cowan, ET. 1929. Life history, habits, and control of
the Mormon Cricket. United States Department of
Agriculture, Technical Bulletin No. 161, 28 pp.
Hitchcock, O. B. 1942. Mormon Cricket Control,
1941—42, pp. 11-13. Jn Mills, H. B., ed., Montana
insect pests, 1941 and 1942. Twenty-ninth report
of the state entomologist. Montana State College
Agricultural Experiment Station Bulletin 408, 36
Pp-
Johnson, N. E 1992. Catalog of world species of Proc-
totrupoidea, exclusive of Platygastridae (Hyme-
noptera). Memoirs of the American Entomologi-
cal Institute 51, 825 pp.
Kozlov, M. A. and S. V. Kononova. 1990. [Scelioni-
nae of the fauna of the USSR: Hymenoptera, Sce-
lionidae, Scelioninae]. Opredeliteli Fauna SSSR
161, 344 pp. [In Russian. ]
Mills, H. B. 1941. Montana insect pests 1939 and
1940. Twenty-eighth report of the state entomol-
ogist. Montana State College Agricultural Exper-
iment Station Bulletin 384, 27 pp.
Muesebeck, C. E W. 1979. Proctotrupoidea, pp.
1121-1186. Jn Krombein, K. V., P. D. Hurd, Jr,
D. R. Smith, B. D. Burks, eds. Catalog of Hy-
menoptera in America North of Mexico. Vol. 1.
Symphyta and Apocrita (Parasitica). Smithsonian
Institution Press, Washington, DC, 1198 pp.
Spencer, G. J. 1958. The natural control complex af-
fecting grasshoppers in the dry belt of British Co-
lumbia. Proceedings of the 10th International
Congress of Entomology (Montreal 1956) 4: 497—
502.
Thorens, P. 1991. Prédateurs et parasites de Chorthip-
pus mollis (Orthoptera, Acrididae) dans deux sta-
tions du pied sud du jura. Bulletin de la Société
Neuchateloise des Sciences Naturelles 114: 43—
Sie
Wakeland, C. 1959. Mormon crickets in North Amer-
ica. United States Department of Agriculture
Technical Bulletin 1202, 77 pp.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 697-704
A REVISION OF NEOTROPICAL DITRICHOPHORA CRESSON
(DIPTERA: EPHYDRIDAE)
WAYNE N. MATHIS
Department of Entomology, NHB 169, Smithsonian Institution, Washington, DC 20560,
U.S.A.
Abstract.—Neotropical species of Ditrichophora Cresson are reported for the first time
from that region and are revised. Two new species are described and illustrated: D. bella
(Dominican Republic. Monsefior Nouel: near Jima (19°01.2’N, 70°28.8’W; 670 m) and
D. chiapas (Mexico. Chiapas: El Triunfo (49 km S Jaltenango)). A diagnosis of the tribe
Discocerinini and an annotated key to New World genera are also provided.
Key Words:
Recent field work on the Dominican Re-
public and Jamaica resulted in the discov-
ery of an undescribed species of Ditricho-
phora Cresson, a genus that has not been
reported from the neotropics (Wirth 1968,
Mathis and Zatwarnicki 1995). The purpose
of this paper is to describe this species,
which is apparently widespread in Central
America and on islands of the West Indies,
and a second Neotropical species that was
recently found in southern Mexico. I am de-
scribing both species within the context of
a revision, including an annotated key to
the New World genera of Discocerinini and
a revised description of the tribe and genus.
Ditrichophora was described in 1924
(Cresson 1924:159) for “‘Discocerine spe-
cies having only two facial bristles. . .”” and
that are “‘... shining and in many respects
resemble those of the Psilopini.’’ The genus
has remained in the tribe Discocerinini
(subfamily Gymnomyzinae) since then
even though the tribe has been recharacter-
ized and now includes only those genera
that are closely related to Discocerina Mac-
quart (Mathis and Zuyin 1989, Mathis and
Zatwarnicki 1995). Other genera that had
been included in the tribe, such as Paratissa
Diptera, Ephydridae, shore flies, Ditrichophora, New World tropics
Coquillett and Rhysophora Cresson, are
now in the tribe Discomyzini (subfamily
Discomyzinae).
Within Discocerinini, Ditrichophora is
closely related and very similar to Gymno-
clasiopa Hendel. Both genera represent bas-
al lineages with Gymnoclasiopa being the
sister lineage to the remaining genera of the
tribe (Zatwarnicki, personal communica-
tion). For many decades, Gymnoclasiopa
was treated as a subgenus within Ditricho-
phora (Cresson 1942, Wirth 1965). Only re-
cently (Zatwarnicki 1992 and personal
communication), with evidence that Gym-
noclasiopa is the most basal lineage in the
tribe, was it recognized as a distinct genus
from Ditrichophora. In this paper, the con-
cept and characterization of Ditrichophora
excludes Gymnoclasiopa, and the latter is
considered a distinct genus in the key to
genera.
Species of Ditrichophora occur through-
out the Old World, but until the discovery
of the two species being described here, the
genus was known only from the Nearctic
Region in the New World. Worldwide, there
are approximately 39 species (Mathis and
Zatwarnicki 1995). Most species occur in
698
temperate, freshwater environments, es-
pecially in the northern hemisphere.
METHODS
The terminology and methods used in
this study were explained previously (Math-
is 1990). Because of the small size of spec-
imens, study and illustration of the male
terminalia required the use of a compound
microscope. To better assure effective com-
munication about structures of the male ter-
minalia, I have adopted the terminology of
other workers in Ephydridae (see references
in Mathis 1986). Usage of these terms,
however, should not be taken as an endorse-
ment of them from a theoretical or morpho-
logical view over alternatives that have
been proposed (Griffiths 1972, McAlpine
1981). Rather, I am deferring to tradition
until the morphological issues are better re-
solved.
Three ratios (one cephalic, two venation-
al) are used commonly in the descriptions
and are defined here for the convenience of
the user (ratios are ranges based on three
specimens).
1. Gena-to-eye ratio is the genal height
measured at the maximum eye height di-
vided by the eye height.
2. Costal vein ratio: the straight line dis-
tance between the apices of veins R,,;
and R,,./distance between the apices of
veins R, and R,,;.
3. M vein ratio: the straight line distance
along M between crossveins dm-cu and
r-m/distance apicad of crossvein dm-cu.
The specimens used in this study are pri-
marily in the National Museum of Natural
History (USNM), Smithsonian Institution.
Paratypes of the Mexican species will be
deposited in the Universidad Nacional Au-
tonoma de México (UNAM).
Tribe Discocerinini Cresson
Discocerinini Cresson, 1925:228. Type ge-
nus: Discocerina Macquart, 1835.—
Mathis and Zuyin, 1989:435 [diagnosis
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
of tribe].—Mathis and Zatwarnicki,
1995:163—186 [world catalog].
Diagnosis.—A tribe of the subfamily
Gymnomyzinae that is distinguished from
other tribes by the following combination
of characters:
Head: Frontal vitta (or ocellar triangle)
mostly bare of setulae, not conspicuously
setulose; ocellar setae well developed, in-
serted anterolaterad of anterior ocellus; rec-
linate fronto-orbital seta inserted antero-
mediad of proclinate fronto-orbital (if 2
proclinate fronto-orbitals present, reclinate
seta inserted anteromediad of the larger,
posterior, proclinate seta); pseudopostocel-
lar setae well developed, proclinate, and
slightly divergent, usually at least half
length of ocellar setae. Pedicel bearing a
large seta anterodorsally; arista bearing 4—
6 dorsal rays, inserted along length of aris-
ta. Face generally shallowly arched, fre-
quently more prominent at level of dorsal
facial setae, not conspicuously pitted, ru-
gose, tuberculate, or carinate. Gena gener-
ally short (secondarily high in some spe-
cies), bearing setulae (including midpor-
tion) and 1 large seta, its posterior (post-
genal) margin rounded, not sharp. Oral
opening and clypeus narrow; mouthparts
generally dark colored.
Thorax: Mesonotum generally microto-
mentose, frequently densely so, although
variable; dorsocentral setae weakly devel-
oped, only posteriormost pair conspicuous;
acrostichal setulae in 2—4 rows, frequently
with a prescutellar pair better developed;
postsutural supra-alar seta usually evident
although sometimes reduced or absent;
prescutellar acrostichal setae inserted ap-
proximate and behind alignment of poster-
iormost dorsocentral setae; scutellar disc
usually densely setulose; scutellum bearing
2 large, marginal setae; notopleural setae 2,
inserted at same level near ventral margin;
anepisternum with 2 subequal setae inserted
along posterior margin. Wing with vein
R,,; long, extended nearly to level of apex
VOLUME 99, NUMBER 4
of vein R,,;. Foreleg normally developed,
not raptorial with greatly enlarged femur.
Abdomen: Male terminalia: Cerci paired,
hemispherical, setose, bearing sides of rec-
tum; epandrium U-shaped, encircling cerci,
anterior margin rounded, in lateral view
with setae mainly on dorsum and along an-
teroventral margin; dististyli lacking or
fused indistinguishably with epandrium;
posterolateral arms of epandrium attached
with ventral apex of gonites, middle of pos-
terior margin a base for aedeagal apodeme;
aedeagal apodeme situated under aedeagus,
associated with hypandrium and with ven-
tral part of base of aedeagus, ventral margin
with lobate appendix providing attachment
for genital muscles that move aedeagus;
gonite paired, connecting sides of base of
aedeagus and laterodorsal margin of epan-
drium, bearing 1 or some setulae; aedeagus
tubular, tapered anteriorly; ejaculatory apo-
deme as a spatula against background of
ductus ejaculatorius.
Discussion.—In our classification for the
family Ephydridae (Mathis and Zatwarnicki
1995), the subfamily Gymnomyzinae La-
treille comprises six tribes, including Dis-
cocerinini. The latter is the most speciose
of the tribes, with 144 of the 346 species
presently included in Gymnomyzinae.
There are eight genera in Discocerinini, and
all eight occur in the New World and are
included in the annotated key that follows.
ANNOTATED KEY TO NEW WORLD GENERA
AND SUBGENERA OF DISCOCERININI
1. Face with secondary series of dorsolaterally in-
clined setae laterad to primary series ......
3 aS cen rete ere ee oer Choe Polytrichophora Cresson
[18 species worldwide; 7 New World species,
presently being revised (Mathis, in prepara-
tion)]
— Face with secondary series of setae lacking or
suggested only by medially inclined setulae .. 2
2. Notopleuron’ bare of setulae .--.......--- 3
— Notopleuron setulose in addition to 2 large se-
PACH etree eRRe CoS OS a3. oh te SS or Shs tens eae Foe Ti
3. Forefemur slightly enlarged, bearing distinct row
of stout, short setae along apical half of postero-
Ventralssunkace es Ame oes oe Pectinifer Cresson
699
[Monotypic; P. aeneus (Cresson), New World
tropics |
Forefemur normally developed, lacking row
of short, stout setae along posteroventral sur-
face fiat Waa wes e Je re ake Sr ee 4
4. Postsutural supra-alar seta strong, distinct, lon-
ger than posterior notopleural seta. Face with
upcurved seta at lower lateral extremity :
cet eA AEE, Che Peay eon = eS Diclasiopa Hendel
[5 species worldwide; a single New World spe-
cies, D. lacteipennis (Loew)]
Postsutural supra-alar seta very short or absent,
if distinguishable distinctly shorter than poste-
rior notopleural seta. Face without upcurved
Setaratelowern:lateralmextnemlity mee eine eee 5)
. Hind tibia with a preapical, ventral, spurlike
seta; facial series comprising 2—3 large setae,
dorsal seta inserted slightly medially from oth-
er setae and arising from distinct, shiny papilla,
with a small, slightly dorsoclinate seta laterad
of dorsal seta; generally microtomentose, ci-
nereous species, appearing dull ..........
SAA = eS ene AE Es Hecamedoides Hendel
[23 species worldwide; a single New World
species, H. unispinosus (Collin)]
Hind tibia lacking a preapical, ventral spurlike
seta; facial series comprised of 2 large setae,
dorsal seta not arising from a shiny papilla and
lacking a smaller seta laterad of dorsal seta;
mostly bare to sparsely microtomentose, shiny
torsubshimysSpeciess i iit enaei eee 6
. Face rather flattened, antennal grooves not al-
ways sharply defined ventrally; facial series of
setae inserted very close to parafacials, dorsal-
most seta not appreciably more removed mesad
thansvenivallsctass ere Gymnoclasiopa Hendel
[25 species worldwide; 8 Nearctic species]
Face rather prominent at level of dorsal facial
setae, sometimes transversely carinate; anten-
nal grooves generally sharply defined ventrally
Li) aye one Sneek a ras Ditrichophora Cresson
[39 species worldwide; 7 Nearctic species, 2
Neotropical species]
. Gena and lower part of parafacial broad; lateral
margin of abdomen usually with gray to whit-
ish microtomentose areas, these usually wedge
shaped. sare eae ee Hydrochasma Hendel
[6 species worldwide, all in the New World]
Gena and parafacial rather narrow; abdomen
lacking wedge-shaped, light-colored areas lat-
erally (genus Discocerina Macquart) ...... 8
[28 species worldwide in 3 subgenera]
= Paratacialabearmousenilac ae ere ac cee
sie fut aes agen subgenus Discocerina Macquart
[9 species worldwide; 7 New World species]
Paratacial lacking setulae “22-2 5.....5-.4-- )
. Facial series of setae 2, these well separated,
distance between subequal to length of Ist fla-
700
gellomere; parafacial very narrow at antero-
ventral margin of eye; postsutural supra-alar
and prescutellar acrostichal setae greatly re-
duced! orlackings>=)5 32) subgenus Basila Cresson
[8 species worldwide, all in the New World]
— Facial series of setae 3—4, distance between
setae conspicuously less than length of Ist fla-
gellomere; parafacial evenly evident through-
out length; postsutural supra-alar and prescu-
tellar acrostichalisetacspresent ye
subgenus Lamproclasiopa Hendel
[11 species worldwide; 9 New World species]
Genus Ditrichophora Cresson
Ditrichophora Cresson, 1924:159. Type
species: Ditrichophora exigua Cresson,
1924, original designation.—Mathis and
Zatwarnicki, 1995:169—174 [world cata-
log].
Strandiscocera Duda, 1942:15. Type spe-
cies: Discocerina nigrithorax Becker,
1926, original designation.—Papp, 1979:
100 [synonymy].
Diagnosis.—Small to medium-sized
shore flies, length 1.25—3.10 mm; generally
mostly bare to sparsely microtomentose,
shiny to subshiny species.
Head: Face rather prominent at level of
dorsal facial seta, sometimes transversely
carinate; antennal grooves generally sharply
defined ventrally; face lacking secondary
series of setae; facial setae usually 2—3, dor-
sal seta not arising from shiny papilla, lack-
ing an upcurved seta at lower lateral ex-
tremity; parafacial narrow throughout
length, lacking setulae; gena generally low.
Eye generally oval, moderately conspicu-
ously microsetulose, bearing several inter-
facetal setulae.
Thorax: Postpronotal and presutural su-
pra-alar setae well developed; postsutural
supra-alar seta reduced or lacking; noto-
pleuron bare of setulae but bearing 2 larger
setae; anterior notopleural seta inserted
closer to posterior notopleural seta than to
postpronotal seta. Forefemur normally de-
veloped, lacking row of short, stout setae
along posteroventral surface; hindtibia lack-
ing a preapical, ventral, spurlike seta.
Abdomen: Abdomen usually unicolor-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ous, lacking wedge shaped, light colored ar-
eas laterally. Fourth tergum of d only
slightly longer than 3rd. Male terminalia as
follows: epandrium complete dorsally, al-
though sometimes attenuated, usually wider
ventrally, especially in lateral view; cercus
hemispherical or elongate (3X as long as
wide), more narrowly pointed dorsally, not
fused anteriorly with epandrium; aedeagus
either simple, mostly tubular, in lateral view
cigar shaped, ovate, or tapered toward apex
or bifurcate apically (best seen in lateral
view) with both lobes large; aedeagal apo-
deme situated behind aedeagus, curved,
keel variously shaped; gonite variously
shaped but generally pointed apically, bear-
ing a subapical or apical setula; ejaculatory
apodeme present, L-shaped; hypandrium
turned up anteriorly, in lateral view irreg-
ularly curved.
Discussion.—The two Neotropical spe-
cies described below are apparently closely
related, both having small, crescent-shaped
indentations on each side of the anterior
portion of the frons above the antennal bas-
es. Moreover, the structures of the male ter-
minalia, especially the external male ter-
minalia (epandrium and cerci), are very
similar.
KEY TO NEOTROPICAL SPECIES OF
DITRICHOPHORA
1. Postpronotum and notopleuron of male gener-
ally bare of microtomentum, shiny, similar to
mesonotum and anepisternum; prescutellar ac-
rostichal setae well developed; male with an-
terior third of frons bare of microtomentum,
shiny black D. chiapas, new species
— Postpronotum and most of notopleuron of male
densely invested with fine, brown microtomen-
tum, contrasted sharply with generally shiny,
adjacent mesonotum and anepisternum; pre-
scutellar acrostichal setae weakly developed:
frons of male generally sparsely microtomen-
tose to anterior margin . D. bella, new species
Ditrichophora bella Mathis, NEW SPECIES
(Figs. 1-3)
Description.—Small to medium-sized
shore flies, length 1.90—3.10 mm; generally
shiny black.
VOLUME 99, NUMBER 4
aedeagal
apodeme
aedeagus
\ postgonite
yA
hypandrium
Figs. 1-4.
701
a
1—3; Ditrichophora bella. 1, External male terminalia (epandrium and surstyli), lateral view. 2,
Same, posterior view. 3, Internal male terminalia (postgonite, pregonite, hypandrium, aedeagal apodeme, aede-
agus), lateral view. 4, Ditrichophora chiapas, Internal male terminalia (postgonite, pregonite, hypandrium, ae-
deagal apodeme, aedeagus), lateral view. Scale = 0.1 mm.
Head: Frons of male generally moder-
ately microtomentose to anterior margin,
only small, linear patch anterolaterally and
at base of setae bare of microtomentum;
frons of female with small bare areas an-
teriorly; 1 proclinate, fronto-orbital seta, in-
serted just behind and laterad of reclinate
seta. Antenna black; apical margin of ped-
icel and Ist flagellomere invested with
whitish gray microtomentum; arista bearing
5 dorsal rays. Face bearing 2 lateral, facial
setae, dorsal seta inserted at level of facial
prominence, ventral seta inserted toward
ventral margin, distance between facial se-
tae about equal to length of Ist flagello-
mere; face black but mostly invested with
silvery white microtomentum, only ventral
portion of antennal grooves, vertical stripe
immediately adjacent to parafacial, and me-
dian stripe that curves laterally ventrally
bare, shiny, black; parafacial completely
microtomentose, whitish gray. Gena-to-eye
ratio 0.1—0.13. Mouthparts, including max-
illary palpus, black.
Thorax: Mesonotum generally sparsely
microtomentose, golden brown, becoming
sparser to bare laterally, through supra-alar
area, thereafter laterally sparsely microto-
702
mentose, male bearing a very distinctive
stripe of dense, fine, brown microtomentum
extended from postpronotum through most
of notopleuron, female shiny black, similar
to adjacent areas; prescutellar acrostichal
setae weakly developed. Wing hyaline; cos-
tal ratio 0.37—0.44; M vein ratio 0.60—0.63.
Legs except tarsi black, mostly shiny, fem-
ora with some surfaces very sparsely mi-
crotomentose; tarsi yellow except apical 2
brown; forefemur unadorned with short,
peglike setulae along posteroventral sur-
face. Halter white.
Abdomen: Black, generally shiny, es-
pecially laterally and ventrally. Male ter-
minalia (Figs. 1—3): Epandrium broadly
rounded dorsally in posterior view (Fig. 2),
narrowed dorsally, becoming wider ventral-
ly, ventral margin bluntly rounded in lateral
view (Fig. 1); cerci lunate in posterior view
(Fig. 2); aedeagus in lateral view slipper-
like, with toe or apex comparatively nar-
rowly rounded (Fig. 3); aedeagal apodeme
more or less triangular in lateral view; post-
gonite and pregonite fused basally; post-
gonite with 2—3 setulae along basoposterior
margin and | setula along margin toward
hypandrium, apex narrowly rounded in lat-
eral view (Fig. 3); hypandrium broadly
rounded along anterior margin, angularly
notched medially with 2 arms attached to
aedeagal apodeme.
Type material.—The holotype ¢ is la-
beled “DOMINICAN RP. Monsf. Nouel:
nr. Jima, 670 m, 19°01.2'N, 70°28.8'WI,]
10 May 1995, W. N. Mathis/HOLOTY PE
Ditrichophora bella ¢ W. N. Mathis USNM
[red; species name and gender handwrit-
ten].””> The holotype is double mounted
(minuten in block of plastic), is in excellent
condition, and is in the USNM. The allo-
type and 24 paratypes (14 6, 10 2; USNM)
bear the same locality label as the holotype.
Other paratypes are as follows: JAMAICA.
St. Andrew: Hardwar Gap (18°04.2'N,
76°44'W), 17 May 1996, D. and W. N.
Mathis, H. Williams (1 6, 1 2; USNM).
MEXICO. Chiapas: El Triunfo (49 km S
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Jaltenango; 1800 m), 14 May 1985, W. N.
Mathis (1 6; USNM).
Distribution.—Neotropical: Mexico (CHI),
West Indies (Dominican Republic, Jamaica).
Diagnosis.—This species is distinguished
from congeners, especially D. chiapas, by
the following combination of characters:
Postpronotum and most of notopleuron of
male densely invested with fine, brown mi-
crotomentum, contrasted sharply with gen-
erally shiny, adjacent mesonotum and an-
episternum; prescutellar acrostichal setae
weakly developed; frons of male generally
sparsely microtomentose to anterior mar-
gin; halter white; only 1 proclinate fronto-
orbital seta; and pattern of silvery white mi-
crotomentum on face (see species descrip-
tion).
Etymology.—The species epithet, bella,
meaning ‘‘pretty, lovely, fine,” is of Latin
derivation and refers to the external attri-
butes of this species.
Remarks.—All specimens were collected
in montane habitats that were frequently
overcast if not enshrouded in a foggy mist.
The specimens from the Dominican Repub-
lic were mostly collected from a pile of
spoiling cabbage that had been discarded on
the roadside.
This species is sexually dimorphic, with
the male have the stripe of dense but fine
microtomentum extended from the post-
pronotum to the posterior margin of the no-
topleuron. Females are shiny black through-
out this area of the pleuron, similar to por-
tions of the mesothorax that are immediate-
ly adjacent.
Ditrichophora chiapas Mathis,
NEw SPECIES
(Fig. 4)
Description.—Moderately small shore
files, length 2.0—2.65 mm; generally shiny
black.
Head: Frons of male and female similar,
moderately microtomentose on posterior
portion, although bare at bases of setulae
and small area just laterad of posterior ocel-
li, anterior third of frons bare, shiny, with
VOLUME 99, NUMBER 4
2 lunate indentations, median area with
pointed extension of moderately microto-
mentum from posterior portion; 1 procli-
nate, fronto-orbital seta, inserted just behind
and laterad of reclinate seta. Antenna black;
apical margin of pedicel and Ist flagello-
mere invested with whitish gray microto-
mentum; arista bearing 5 dorsal rays. Face
bearing 3 lateral, facial setae, dorsal seta
longest, inserted at level of facial promi-
nence, ventral setae progressively shorter,
evenly spaced with distance between less
than width of Ist flagellomere; face mostly
black, shiny, but with pattern of silvery
white microtomentose, vertical stripes; lat-
eral facial stripe immediately adjacent to
parafacial, other vertical stripe just laterad
of midfacial vertical bare area; also silvery
white microtomentose on dorsal portion of
antennal grooves and along ventral, facial
margin; parafacial bare, shiny black. Gena-
to-eye ratio 0.1—0.12. Mouthparts, includ-
ing maxillary palpus, black.
Thorax: Mesonotum generally sparsely
microtomentose, golden brown, becoming
bare laterally through supra-alar area and
continuing ventral through pleural area;
prescutellar acrostichal setae well devel-
oped. Wing hyaline; costal ratio 0.40—0.43;
M vein ratio 0.70—0.73. Legs except tarsi
black, mostly shiny, femora with some sur-
faces very sparsely microtomentose; tarsi
yellow except apical 1—2 brown; forefemur
with row of numerous, very short, peglike
setulae along posteroventral surface. Halter
white.
Abdomen: Black, generally shiny, es-
pecially laterally and ventrally. Male ter-
minalia (Fig. 4): Epandrium and cerci as in
D. bella; aedeagus in lateral view slipper-
like, with toe or apex comparatively bluntly
rounded (Fig. 4); aedeagal apodeme more
or less triangular in lateral view (Fig. 4);
postgonite and pregonite fused basally;
postgonite with | setula along basoposterior
margin and | setula along margin toward
hypandrium, apex acutely rounded in lateral
view (Fig. 4); hypandrium broadly rounded
along anterior margin, angularly notched
703
medially with 2 arms attached to aedeagal
apodeme.
Type material—The holotype ¢ is la-
beled ““MEXICO. Chiapas: El Triunfo (49
km S Jaltenango) 14 May 1985, 1800 m[,]
Wayne N. Mathis/HOLOTYPE Ditricho-
phora chiapas W. N. Mathis USNM [red;
species name handwritten].’’ The holotype
is double mounted (minuten in block of
plastic), is in excellent condition, and is in
the USNM. The allotype 2 and four para-
types (4 6; USNM) bear the same locality
label as the holotype. Other paratypes are
as follows: MEXICO. Chiapas: El Triunfo
(49 km S Jaltenango; 1300-2000 m), 13-
15 May 1985, W. N. Mathis (3 6; UNAM,
USNM).
Distribution.—Neotropical:
(CHI).
Diagnosis.—This species is distinguished
from congeners, especially D. bella, by the
following combination of characters: Post-
pronotum and notopleuron of male gener-
ally bare of microtomentum, shiny, similar
to mesonotum and anepisternum; prescutel-
lar acrostichal setae well developed; male
with anterior third of frons bare of micro-
tomentum, shiny black; halter white; only 1
proclinate fronto-orbital seta; and pattern of
silvery white microtomentum on face (see
species description).
Etymology.—The species epithet, chia-
pas, refers to the Mexican state where the
type series was collected and is treated as a
noun in apposition.
Remarks.—El Triunfo is a site in the
cloud forest of southern Mexico (some of
the only cloud forest that remains largely
undisturbed in Mexico).
Mexico
ACKNOWLEDGMENTS
I am grateful for assistance in the field
from Oliver S. Flint, Jr. (USNM), Amnon
Freidberg (Tel-Aviv University), Kelvin A.
Guerrero (Museo Nacional de Historia Nat-
ural, Santo Domingo), N. Dianne Mathis
(USNM), and Hollis B. Williams (USNM).
For critically reviewing a draft of this paper,
I thank Volker Hollmann-Schirrmacher and
704
Allen L. Norrbom. The illustrations were
skillfully inked by Elaine R. S. Hodges.
Tadeusz Zatwarnicki (Department of Zool-
ogy, University of Agriculture, ul. Cybul-
skiego 20, 50-205 Wroclaw, Poland) and I
are working on a generic revision of Dis-
cocerinini, and our joint research and many
discussions have contributed to my under-
standing of the tribe and this revision.
Funding for this research project, es-
pecially the field work on the Dominican
Republic, was provided by the Biodiversity
Program (Biological Surveys and Invento-
ries, BSI), National Museum of Natural
History, Smithsonian Institution (Lynne R.
Parenti, chair).
LITERATURE CITED
Cresson, E. T., Jr. 1924. Descriptions of new genera
and species of the dipterous family Ephydridae.
Paper VI. Entomological News 35(5): 159-164.
. 1925. Descriptions of new genera and species
of the dipterous family Ephydridae. VII. Ento-
mological News 36(6): 165-167.
1942. Synopses of North American Ephyd-
ridae (Diptera) I. The Subfamily Psilopinae, with
Descriptions of New Species. Transactions of the
American Entomological Society 68: 101-128.
Duda, O. 1942. Neue oder ungeniigend bekannte
Zweifliigler der palaéarktischen Region aus meiner
Sammlung. 2. Fortsetzung. Deutsche Entomolo-
gische Zeitschrift 1-4: 1—39.
Griffiths, G. C. D. 1972. The Phylogenetic Classifi-
cation of Diptera Cyclorrhapha with Special Ref-
erence to the Structure of the Male Postabdomen.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Series Entomologica 8:
Hague.
Mathis, W. N. 1986. Studies of Psilopinae (Diptera:
Ephydridae), I: A revision of the shore fly genus
Placopsidella Kertész. Smithsonian Contributions
to Zoology 430: 1-30.
1990. A revision of the shore-fly genus Di-
phuia Cresson (Diptera: Ephydridae). Proceedings
of the Entomological Society of Washington
92(4): 746-756.
Mathis, W. N. and T. Zatwarnicki. 1995. A world cat-
alog of the shore flies (Diptera: Ephydridae).
Memoirs on Entomology, International 4: vi +
423 pp.
Mathis, W. N. and J. Zuyin. 1989. A review of the
shore-fly genus Polytrichophora Cresson from
Asia (Diptera: Ephydridae). Proceedings of the
Biological Society of Washington 102(2): 434—
446.
McAlpine, J. EF 1981. Morphology and terminology-
adults, pp. 9-63. In McAlpine, J. F, et al., eds.,
Manual of Nearctic Diptera. Ottawa. [Volume | is
Monograph 27 of Research Branch Agriculture
Canada. |
Papp, L. 1979. A contribution to the revision of the
Palaearctic Ephydridae (Diptera). Folia Entomo-
logica Hungarica, series nova 32(1): 97—104.
Wirth, W. W. 1965. Ephydridae, pp. 734-759. In
Stone, A., C. W. Sabrosky, W. W. Wirth, R. H.
Foote, and J. R. Coulson, eds., A Catalog of the
Diptera North of Mexico. Handbook, 276: 1696
pp. U.S. Department of Agriculture, Washington,
IDE
1-340 pp. W. Junk,
1968. 77. Family Ephydridae, pp. 1—43. In
Papavero, N., ed., A catalogue of the Diptera of
the Americas south of the United States. Depar-
tamento de Zoologia, Secretaria de Agricultura,
Sao Paulo.
Zatwarnicki, T. 1992. A new classification of Ephyd-
ridae based on phylogenetic reconstruction (Dip-
tera: Cyclorrhapha). Genus 3(2): 65-119.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 705-719
NEW SPECIES AND HOST PLANTS OF THE ANTHONOMUS GRANDIS
SPECIES GROUP (COLEOPTERA: CURCULIONIDAE)
ROBERT W. JONES AND HORACE R. BURKE
(RWJ) El Colegio de la Frontera Sur, Carretera Panamerica y Periférico Sur, Apartado
Postal 63, San Cristobal de las Casas, Chiapas, México (present address: Licenciatura en
Biologia, Universidad Aut6énoma de Querétaro, AP184, Querétaro, QRO, México
CP76010); (HRB) Department of Entomology, Texas A&M University, College Station,
TX 77843-2475, U.S.A.
Abstract.—Three new species of the Anthonomus grandis group are described and
illustrated, and a key distinguishing the five species of the group is presented.
Anthonomus townsendi, n. sp., and A. palmeri, n. sp., occur in the State of Chiapas,
México, and A. mallyi, n. sp., occurs in Costa Rica. Host plants of these species are
members of the genus Hampea (Malvales: Malvaceae). The taxonomic and ecological
interrelationships of the weevil taxa and their malvaceous hosts are discussed.
Key Words:
hosts
Since the boll weevil, Anthonomus gran-
dis Boheman, was first recognized as a pest
of cotton in the late 1800’s considerable ef-
fort has been expended in attempts to un-
derstand its origin, systematics, and host re-
lationships (Burke et al. 1986). Until re-
cently, the lack of knowledge of relatives
of the boll weevil hindered meaningful
comparative studies of its phylogeny and
ecology. Anthonomus hunteri Burke and
Cate, the first species recognized as having
close morphological affinities to A. grandis,
was described in 1979 (Burke and Cate
1979). Before the present study, these were
the only two known species of the A. gran-
dis group as defined by Clark and Burke
(1986). Three new species are added to the
group in the present paper.
Both of the previously known species of
this group are almost entirely restricted to
larval development on certain Malvaceae of
the tribe Gossypieae, including Gossypium,
Hampea, Cienfuegosia, and Thespesia. An-
Curculionidae, Anthonomus, new species, México, Costa Rica, Hampea
thonomus hunteri is known to develop only
on Hampea trilobata Standley, a plant en-
demic to the Yucatan Peninsula, Belize and
northern Guatemala, while A. grandis has a
wider host range within Gossypieae (Burke
et al. 1986; Grosset al. 1975, Pry xell-and
Lukefahr 1967, Fryxell 1969). Anthonomus
grandis was thought to be specific to Gos-
sypieae until Bodegas et al. (1977) discov-
ered members of the species developing on
Hibiscus pernambucensis Arruda of the
tribe Hibisceae in Chiapas, México.
A growing body of evidence indicates
that Hampea is the ancestral host plant ge-
nus of the A. grandis species group. Both
previously known species of the group uti-
lize Hampea as hosts, and A. hunteri de-
velops exclusively on H. trilobata. The
three new species of the A. grandis group
described herein also utilize species of
Hampea as hosts. Further evidence of an
ancestral relationship of the weevils with
plants of the genus Hampea includes the
706
following: 1) large populations of A. gran-
dis occur on Hampea nutricia Fryxell in
southern México in the absence of cultivat-
ed or wild cotton; 2) the parasite guild of
weevils on Hampea is more diverse than
that for weevil populations on cotton
(Burke et al. 1986, Cate et al. 1990); and,
3) A. grandis and A. hunteri have behav-
ioral adaptations apparently specific to
Hampea plants that suggest long periods of
association (Stansly 1985).
Since Hampea has been shown to be the
likely original host genus of the boll weevil,
considerable interest has been generated re-
garding the ecology of these plants and
their interactions with weevils. Hampea
contains 21 described species from both the
Gulf and Pacific coasts of México and
southward into Colombia, with the greatest
species diversity occurring in Chiapas,
México and Guatemala (Fryxell 1969,
1979). Only four species of Hampea have
been previously reported as hosts of An-
thonomus, three of these being utilized by
A. grandis and one by A. hunteri. Nine ad-
ditional species of Hampea were examined
during the present study. Four of these were
found to serve as hosts of three heretofore
undescribed species of the A. grandis
group. With the A. grandis species group
now being comprised of five species, it is
possible to conduct a detailed comparison
of the these taxa to assist in analyzing their
origin, ecology, and host and phylogenetic
relationships. The objectives of the present
paper are to describe the three new species,
compare them with their previously known
relatives, and to report on the current status
of eight species of Hampea in México and
Central America as hosts of members of the
A. grandis species group. A cladistic anal-
ysis of both the weevils and their Hampea
hosts is underway. Additional papers on the
ecology and genetics of this weevil group
are also being prepared.
METHODS AND MATERIALS
Determination of the status of species of
Hampea as hosts of Anthonomus.—During
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
the years 1987-1990, 1992, and 1995 nine
species of Hampea not previously known
to be hosts of Anthonomus were examined
for evidence of weevil infestations in Mé-
xico, Guatemala, Belize, and Costa Rica.
Examined were: Hampea appendiculata (J.
Donnell-Smith) Standley, Hampea bracteo-
lata Lundell, Hampea integerrima Schlech-
tendal, Hampea longipes Miranda, Hampea
mexicana Fryxell, Hampea montebellensis
Fryxell, Hampea platanifolia Standley,
Hampea stipitata S. Watson, and Hampea
tomentosa (K. Presl) Standley. These spe-
cies were located in the field on the basis
of distributional records obtained from
specimens in the following herbaria: United
States Department of Agriculture Cotton
Laboratory, College Station, TX; Herbario
Nacional, Universidad Aut6noma de Méx-
ico, México, D.E; and University of Texas,
Austin, TX. Paul A. Fryxell (pers. comm.),
specialist on the systematics of this group,
also provided additional information useful
in locating the plants. Each tree was ex-
amined for the presence of weevils and for
evidence of feeding and oviposition both on
the tree and in fallen flower buds. Poten-
tially infested buds were enclosed in vials
in the laboratory to allow for emergence of
adult weevils.
Comparisons and descriptions of wee-
vils.—To establish the relationships of the
weevils collected and reared during this
study, it was necessary to compare speci-
mens reared from Hampea with those from
other malvaceous hosts. A total of 761
adults of the A. grandis species group from
13 species of Malvaceae was examined. In
addition to weevils from eight species of
Hampea, material was studied from Gos-
sypium hirsutum L., Gossypium thurberi
Todaro, Cienfuegosia rosei Fryxell, Cien-
fuegosia drummondii A. Gray, and Hibiscus
pernambucensis Arruda.
Comparisons were made between adult
weevils from the entire range of known
hosts plants. Characters used by Burke
(1986) and Burke et al. (1986) to distin-
guish the three recognizable forms of A.
VOLUME 99, NUMBER 4
grandis (Southeastern, Mexican, and Thur-
beria), and by Clark and Burke (1986) in a
phylogenetic study of the A. grandis group
were used here. Characters examined in-
cluded the shape and arrangement of pro-
notal and elytral scales, shape of profemur,
shape and alignment of scutellum, colora-
tion of antennae, and size and shape of
body. The male median lobe proved to be
especially useful in distinguishing species
of the group. Analysis of specimens of A.
grandis reared from various host plants aid-
ed in evaluation of the effects of these hosts
on the general morphology and intraspecific
variation within the species group. A con-
siderable amount of previously accumulat-
ed information on host-induced and geo-
graphic variation of A. grandis provided a
basis for comparison and evaluation of the
taxonomic status of the newly collected
weevils on Hampea.
Measurements were made with an eye-
piece micrometer in a stereoscopic micro-
scope. The size of the weevils was deter-
mined in a lateral view by measuring from
the anterior margin of the eye to the apex
of the elytra. The length of the rostrum was
determined by measuring the chord from
the lower anterior margin of the eye to the
apex of the rostrum. Other structures were
measured at the point of their greatest
length or width.
RESULTS
Host plants.—Weevils of the A. grandis
group were reared from four species of
Hampea not previously reported as hosts:
H. appendiculata (Coastal Plain of eastern
Costa Rica); H. longipes (Central High-
lands of Chiapas); H. mexicana (Central
Depression of Chiapas); and H. montebel-
lensis (eastern Chiapas). Five additional
species of Hampea (H. bracteolata, H. in-
tegerrima, H. platanifolia, H. stipitata, and
H. tomentosa) were examined but no wee-
vils were found associated with these. Sev-
eral relatively large populations of H. sti-
piata and H. integerrima proved to be un-
infested during a period of two years of
707
monitoring. Both of these species of plants
were heavily laden with flower buds at the
times of observation and, furthermore, they
occurred within the flight range of A. gran-
dis on H. nutricia in the states of Veracruz
and Chiapas. Conditions appeared to be ex-
cellent for weevils to develop on these
plants but accumulated evidence indicated
that they did not do so. Because of their
isolated distributions and relative rarity,
only small populations of H. platanifolia,
H. bracteolata, and H. tomentosa were
found. Although the few trees observed
were large and bore numerous flower buds,
they were apparently not infested with wee-
vils.
Definition of group.—Anthonomus gran-
dis, A. hunteri and the three new species
described herein are considered to be mem-
bers of the same species group because of
their close morphological and host plant af-
finities, although it is not possible to de-
scribe the group precisely on the basis of
shared apomorphies. All of the species of
the group develop in flower buds, and oc-
casionally fruits, of Malvaceae of the tribe
Gossypieae. In addition to its cotton tribe
hosts, one population of A. grandis devel-
ops on Hibiscus pernambucensis, tribe Hi-
bisceae. The latter association is apparently
the result of a recent, localized host shift
which should not detract from the fact that
the members of the group obviously
evolved on closely related hosts in the tribe
Gossypieae, probably in the genus Hampea.
The weevils of this group are also of a rel-
atively large size for the genus, generally
averaging approximately 4 to 5 mm in
length and are occasionally larger. There are
other tropical anthonomines of similar or
larger size, for example in the genus Lon-
cophorus, but the relationship of these to
Anthonomus, and specifically to the A.
grandis group, has not been determined
(see Clark 1988, 1995, for further discus-
sion of this problem). Furthermore, known
hosts of species of Loncophorus are all
members of the families Bombacaceae and
Tiliaceae. While these plant families are
708 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
closely related to Malvaceae we do not be-
lieve that this, in itself, is sufficient evi-
dence to closely ally Lonchophorus and the
A. grandis group. In addition to the char-
acters mentioned above that are shared by
species of the A. grandis group, they have
similar subconical heads, coarse hair-like to
scale-like setae that are more-or-less evenly
distributed over the body, rounded sides of
the elytra as compared with the wedge-
shaped or “‘navicular’’ elytra characteristic
of most Lonchophorus, and do not have
prothoracic postocular lobes as do species
of the latter genus. The A. grandis group
was compared to the subgenus Anthono-
morphus Dietz of the genus Anthonomus by
Clark and Burke (1986). Species of An-
thonomorphus also develop on malvaceous
plants, but unlike the A. grandis group,
their hosts are in the tribe Malveae. In ad-
dition, the two groups differ in several mor-
phological characters, most notably in the
shapes of the male mesotrochanters and
median lobes. While Anthonomorphus and
the A. grandis species group may not be
sister groups, there is still sufficient evi-
dence to warrant further consideration of
their relationships. Clark (1995) suggested
that Lonchophorus, Anthonomorphus, and
the A. grandis species group may constitute
a monophyletic lineage confined to the
Malvales. Additional study of certain Neo-
tropical members of the tribe Anthonomini,
especially the species of Lonchophorus and
its allies, is needed to help clarify the over-
all relationships of species of the A. grandis
group. Pending such a study, size, general
habitus, vestiture, and narrow host ranges
restricted almost entirely to plants in the
tribe Gossypieae are used here to define the
group.
KEY TO SPECIES
1. Rostrum slightly to moderately strongly cari-
nate dorsally; hirsute only at base or may be
nearly to completely bare; female rostrum
slightly to moderately strongly curved (Figs. 3,
4, 5, 6). Elytral vestiture usually arranged in an
orderly manner, with individual setae directed
i)
posteriorly. Male median lobe moderately to
strongly curved in lateral view (Figs. 23, 24,
26, 27). Protibia slightly to moderately strongly
Sinnate Oni immer ar oes ee 2,
Rostrum strongly carinate and hirsute dorsally
from base to antennal attachments; female ros-
trum nearly straight (Fig. 1). Elytral vestiture
arranged in irregular manner, with individual
setae often directed obliquely or laterally. Male
median lobe broadly rounded at apex; slightly
curved ventrally in lateral view (Fig. 25). Pro-
tibia strongly sinuate on inner margin. Hosts
Hampea mexicana and H. latifolia. Central and
southern Chiapas, México, above 500m....
Sis ison tones eae A. townsendi Jones and Burke, n.sp.
. Rostrum of female slender; dorsal carinae
poorly defined. Antennae of female attached at
or near middle of rostrum. Male median lobe
slightly (Fig. 24) to strongly (Fig. 26) constrict-
ed before tipo hae. ee eee 3
Rostrum of female stouter; dorsal carinae mod-
erately strongly developed. Antennae of female
usually attached well before middle of rostrum.
Male median lobe broadly rounded to some-
what pointed at tip, but sides not constricted
(Figs:[23 27) = Sos eae ea 4
. Rostrum of both sexes moderately strongly
curved. Profemur slender, length/width ratio
4.2—4.7. Male median lobe with sides slightly
constricted before apex (Fig. 24). Host Ham-
pea trilobata. Yucatan Peninsula, México, and
Belizet J 334 Ai See A. hunteri Burke and Cate
Rostrum of both sexes slightly curved (Figs. 3,
4). Profemur stouter, length/width ratio 3.0—
3.5. Male median lobe strongly constricted be-
fore apex (Fig. 26). Host Hampea appendicu-
lata. ‘Costa "Rica “2c. Acs) ees ee
aL Monee Crees A. mallyi Jones and Burke, n. sp.
. Median vitta of setae on pronotum well de-
fined. Setae on procoxae and sides of prothorax
scale-like, with blunt apices (Figs. 18, 22).
Male median lobe moderately strongly curved
in lateral view; sides of apical portion con-
verging strongly to blunt tip; length/width ra-
tios of apical portion (measured from retracted
phallotreme to tip) 1.2-1.7 (Fig. 23). Hosts
Gossypium spp., Cienfuegosia spp., Thespesia
populnea, Hibiscus pernambucensis, Hampea
nutricia, H. rovirosae, and H. latifolia. United
States, México, Central America, Venezuela,
Colombia, Brazil, and Argentina .........
60s GG EE ioe eo eee A. grandis Boheman
Median vitta of setae on pronotum absent or
poorly defined; setae on procoxae and sides of
prothorax narrow, with apices sharply pointed
(Figs. 17, 21). Male median lobe slightly
curved in lateral view; sides of apical portion
broadly rounded to tip; length/width ratios of
VOLUME 99, NUMBER 4
Figs. 1-6.
1, Anthonomus townsendi, female. 2, A. townsendi, male. 3, Anthonomus mallyi, female. 4, A.
709
mallyi, male. 5, Anthonomus palmeri, female. 6, A. palmeri, male.
apical portion 0.7—1.0 (Fig. 27). Hosts Hampea
montebellensis and H. longipes. Central High-
lands and eastern Chiapas, México, above
LOOOhnae A. palmeri Jones and Burke, n. sp.
SPECIES DESCRIPTIONS
Anthonomus townsendi Jones and
Burke, NEW SPECIES
(ncaa 9258s. e aL Oe 25)
Body oblong-oval; length 4.17—6.25 mm
(x = 5.62, n = 30); width 1.92—2.92 mm
(x = 2.60, n = 30). Integument shining,
dark brown; head and rostrum sometimes
darker than thorax and abdomen. Vestiture
of coarse, recumbent, straw-colored pubes-
cence; arranged in irregular patches on el-
ytra (Figs. 1, 2); concentrated along midline
of pronotum; elsewhere on prothorax fine
and slightly broader setae intermixed; on
underside and legs pubescence uniformly
distributed. Rostrum of both sexes feebly
curved, that of male (Fig. 2) being slightly
more strongly curved than female (Fig. 1);
ca. 2X longer than prothorax along midline
710
in both sexes. Median and lateral rostral ca-
rinae strongly developed in male, less so in
female; carinae and associated pubescence
of male extending from base past antennal
insertion (Fig. 7), in female extending only
to level of antennal insertion (Fig. 8). Dis-
tad of antennal insertion rostrum of male
with deep, oval punctures, each bearing a
seta longer than greatest diameter of punc-
ture (Fig. 7), rostrum of female not as
strongly punctate distally (Fig. 8). Eyes
moderately convex in dorsal view; not free
behind; separated in front by distance equal
to ¥; width of rostrum at base. Frons deeply
foveate; moderately densely covered with
fine coarse pubescence. Antennae attached
ca. apical % of rostrum of male, ca. apical
¥% in female. Funicle 7-segmented, 1.7—
2.2 longer than club; segment 1 slender,
as long as next two segments combined;
segment 2 slightly longer than 3 + 4. Club
subequal to preceding five funicular seg-
ments combined; basal two segments loose-
ly joined; apex sharply pointed. Prothorax
with sides feebly rounded, subparallel in
basal % then converging to subapical con-
striction; width at apex % width at base; in-
tegument shining, bearing large, rounded
punctures that are smaller medially than lat-
erally, areas between punctures smooth,
bare, each puncture bearing an anteriorly
projecting seta; vestiture concentrated into
moderately dense median pronotal vitta,
uniformly distributed laterally (Fig. 15) ex-
cept slightly concentrated above coxae.
Scutellum ca. 2X longer than wide, rounded
in cross section, sides parallel in apical %,
expanding slightly at base; clothed above
with appressed, fine scale-like setae that are
concentrated in basal % and densely clothed
at apex. Elytra strongly convex (Figs. 1, 2)
in lateral view; slightly wider at base than
prothorax; humeri moderately strongly
rounded; sides subparallel to point past
middle then strongly rounded to apex.
Third elytral interval sometimes expanded
basally. Pubescence arranged in regularly
spaced patches presenting marmorate ap-
pearance, sometimes coalescing into more
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
linear patterns towards elytral apex. Inter-
vals fairly strongly convex, especially on
basal % of elytra. Ventral side of body with
pubescence on thorax coarse, more hair-like
on abdominal sterna. Procoxae contiguous.
Mesocoxae separated by distance equal to
ca. % width of coxa. Bare depression me-
dially near posterior margin of metaster-
num. Abdominal sternum 1 along midline
slightly longer than 2; sternum 3 ca. % lon-
ger than 4. Legs moderately stout (Figs. 1,
2); rather uniformly covered with coarse
pubescence. Profemur 3.2—3.6X longer than
broad; bearing two teeth, the proximal of
which is distinctly larger than triangular
distal tooth. Meso- and metafemur ca. equal
in size, each bearing a small tooth. Protibia
curved in basal % and sinuate on inner mar-
gin (Fig. 1); meso- and metatibia straight
and less strongly sinuate on inner margins.
Tarsal claws each bearing long, slender
tooth, the apex of which reaches past mid-
dle of claw. Median lobe of male genitalia
feebly curved in lateral view; apex broadly
rounded, distance from retracted phailotre-
me to apex of median lobe less than width
of median lobe (Fig. 25).
Type series.—Holotype d-México: 30
km N. D. Chanona, Chiapas, VI-29-1990,
ex. Hampea mexicana (buds on ground), R.
Jones. Paratypes (68): 9 3d, 8 2 with same
label data as holotype. México: 9 km N.
Nvo. Tenochititlan, Mpio. Cintalapa, Chia-
pas, ex. Hampea mexicana, R. W. Jones (1
2, VII-23-1988; 4 6, 1 2, VI-12-1991; 2
3, 2 2, VI-28-1991; 6 6, 4 2, VH-12-
1991). México: 5 mi. N. Nvo. Tenochititlan,
Mpio. Cintalapa, Chiapas, VI-29-1990, ex.
Hampea mexicana, R. W. Jones and D.
Baro (11 6, 11 2). México: Jiquipilas, 4
mi W. Fco. Villa, Chiapas, VIII-6-1989,
Jones, Cate and Krauter, ex. Hampea mex-
icana (1 6, 3 2). México: 19 km N. Ma-
pastepec, Chiapas, VII-7, 1991, R. Jones,
ex. Hampea latifolia ground buds (2 d, 1
2); 1 3, 1 2 same data as preceding except
VITI-11-1991.
The holotype and some paratypes are de-
posited in the Insect Collection of the De-
VOLUME 99, NUMBER 4
711
Figs. 7-14. Rostra of: 7, Anthonomus townsendi, male. 8, A. townsendi, female. 9, Anthonomus palmeri,
male. 10, A. palmeri, female. 11, Anthonomus mallyi, male. 12, A. mallyi, female. 13, Anthonomus grandis,
male. 14, A. grandis, female.
partment of Entomology, Texas A&M Uni-
versity. Paratypes are deposited in the col-
lections of: Instituto de Biologia, Univer-
sidad Aut6noma de México, D.F; El
Colegio de la Frontera Sur, San Cristobal
de las Casas, Chiapas, México; Auburn
University; Canadian Museum of Nature,
Ottawa, Canada; and Charles W. O’Brien.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 15-22. Ventral one-half of pleural region of prothorax of: 15, Anthonomus townsendi. 16, Anthonomus
mallyi. 17, Anthonomus palmeri. 18, Anthonomus grandis (ex Hampea nutricia). Dorsal one-half of pleural
region of prothorax (above base of procoxa) of: 19, Anthonomus townsendi. 20, Anthonomus mallyi. 21, An-
thonomus palmeri. 22, Anthonomus grandis (ex Hampea nutricia).
Diagnosis and variation.—Characters
differentiating species of the A. grandis
group are presented in the key. In general,
A. townsendi is larger and more robust than
other species of the group. The marmorate
appearance of the pubesence on the elytra
is also distinctive. The female rostrum is
less strongly curved than in the other spe-
cies and the apical length (from antennal
attachment to apex of rostrum) is distinctly
VOLUME 99, NUMBER 4
Figs. 23-27.
shorter relative to the rostrum width. The
male rostrum of A. townsendi is the most
densely hirsute of the members of the com-
plex and also has the most strongly pro-
nounced punctures and largest scales distad
of the antennal insertion (Fig. 7). The pro-
tibia of A. townsendi is more strongly
curved and the inner margin is more strong-
ly sinuate than that of either A. grandis, A.
hunteri or A. mallyi. The protibia of A. pal-
meri is curved basally like that of A. town-
sendi but as in other members of the group
it is slender and the inner margin is not as
sinuate. Furthermore, A. townsendi may be
distinguished from the three recognized
forms of A. grandis by the pubescence be-
ing uniformly distributed on the sides of the
prothorax. The relatively straight median
lobe in lateral view with the apex being
wider than long also distinguishes this spe-
cies from A. grandis, A. hunteri, and A.
mallyi.
The major variation in the type series is
in size. Specimens range from 4.2 to 6.3
Dorsal and lateral views of male median lobe of: 23, Anthonomus grandis (ex Hampea nutricia).
24, Anthonomus hunteri. 25, Anthonomus townsendi. 26, Anthonomus mallyi. 27, Anthonomus palmeri.
mm in length. Teeth on the profemur show
considerable variation in relative size. On
some individuals the profemoral teeth are
similar in size and shape to those of A.
grandis. However, the smaller of the two
profemoral teeth of A. townsendi is some-
times strongly reduced in comparison with
the larger one. The larger profemoral tooth
of A. townsendi is broader at the base than
that of A. grandis. As is often the case with
reared material, some of the individuals of
the type series are somewhat teneral and are
thus not as dark as would be expected of
well hardened specimens.
Host plants.—Hosts of this species are H.
mexicana in the Central Depression area of
Chiapas, and H. latifolia on the slopes of
the Sierra Madre de Chiapas above 500 m
elevation. It is of interest to note that H.
latifolia is also a host of A. grandis on the
coastal plain near Tapachula, Chiapas.
Specimens reared from the latter host pop-
ulation were clearly A. grandis and showed
no overlap in distinguishing characters with
714
A. townsendi reared from H. latifolia in
montane habitats of the Sierra Madre de
Chiapas, approximately 100 km east of the
A. grandis population. It is unknown
whether the utilization of the same species
of Hampea by the two weevils is the result
of different habitat preferences or the result
of a relatively recent displacement of A.
townsendi by enormous populations of A.
grandis developing on cultivated cotton in
the coastal plain.
Derivation of name.—This species is
named in honor of Charles Henry Tyler
Townsend (1863-1944), the first entomol-
ogist to conduct scientific study of the boll
weevil. Townsend studied the boll weevil
under difficult conditions in South Texas in
the late 1800’s immediately after the first
discovery of the pest in the United States
(Townsend 1895). He also traveled widely
in México and made extensive collections
of insects there.
Anthonomus mallyi Jones and Burke,
NEW SPECIES
(Pigss 3.24 S112 16, 20,26)
Body elongate-oval; length 4.67—5.17
mm (x = 4.98, n = 7); width 2.17—2.42 mm
(x = 2.32, n = 7). Integument shining, tan,
head and rostrum brown (type series reared;
specimens somewhat teneral; well hardened
specimens expected to be darker). Vestiture
sparse, consisting of fine, yellowish white,
prostrate pubescence; shorter and irregular-
ly distributed in patches on elytra (Figs. 3,
4); faint median vitta on pronotum; no-
where on body completely obscuring integ-
ument. Rostrum slightly curved in both sex-
es, that of female being nearly straight
(Figs. 3, 4); rostrum of male 1.8 and that
of female 2.5 longer than prothorax along
dorsal midline; ratios of rostrum length past
antennal insertion to width (measured in
side view) 1.7 for male and 4.0 for female.
Pubescence on rostrum of both sexes
sparse, inconspicuous except near base, not
extending distad of level of antennal inser-
tions (Figs. 11, 12). Median and lateral ca-
rinae rounded, with intervening grooves
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
shallow, not quite reaching level of anten-
nal insertions. Lateral rostral groove rela-
tively well defined; upper margin imping-
ing against front margin eye at about mid-
dle. Eyes moderately strongly convex in
dorsal view, not free behind; separated in
front by distance equal to ca. % width of
rostrum at base. Frons foveate; vestiture
sparse, about same density as on pronotum.
Antennae attached at ca. apical %4 of ros-
trum in male and slightly distad of middle
in female. Funicle 7-segmented; 1.5—1.8X
longer than club; segment 1 slender, 2X
length of segment 2, ca. as long as follow-
ing four segments combined; remaining
segments ca. equal in length. Club elongate,
ca. as long as preceding six funicular seg-
ments combined; basal two segments loose-
ly joined; apex sharply pointed. Prothorax
with sides parallel in basal % then converg-
ing moderately strongly to subapical con-
striction; integument shining; elongate
punctures smaller dorsally than laterally,
each bearing an anteriorly pointing seta; ar-
eas between punctures bare, impunctate.
Vestiture coalescing into faint to relatively
well defined median vitta, laterally on pro-
notum uniformily distributed; scale-like se-
tae on pleural region of prothorax striate
(Fig. 20), stouter than those dorsally. Scu-
tellum rectangular, parallel-sided, somewhat
flattened dorsally, about 1.5 longer than
wide; sparsely clothed with appressed, fine
setae. Elytra in lateral view moderately
strongly convex at base in dorsal % (Figs.
3, 4); distinctly wider at base than protho-
rax; humeri moderately strongly rounded;
sides subparallel to point past middle then
strongly rounded to apex. Basal margin
from third interval outward relatively
straight. Pubescence arranged in more-or-
less regularly spaced patches on basal % of
elytra, coalescing into more linear patterns
towards elytral apex. Intervals somewhat
flattened. Ventral side of body clothed with
pubescence that does not completely ob-
scure integument, finer and less dense on
legs. Procoxae contiguous. Mesocoxae sep-
arated by distance equal to ca. % width of
VOLUME 99, NUMBER 4
coxa. Abdominal sternum 1 along midline
equal in length to sterna 2 + 3. Legs mod-
erately stout (Figs. 3, 4). Profemora 3.0—
3.5 longer than wide; bearing two teeth;
basal tooth slightly quadrate, 1.8—3.1X lon-
ger than broadly triangular distal tooth.
Meso- and metafemora ca. equal in size,
each bearing a small, sharply pointed tooth.
Protibia relatively straight, moderately sin-
uate on inner margin. Meso- and metatibiae
each slightly sinuate on inner margin. Tar-
sal claws each bearing a long tooth on inner
margin, the apex or which reaches past the
middle of claw. Median lobe of male
strongly curved in lateral view, especially
in distal %, length of apex (distance from
retracted phallotreme to apex of lobe) great-
er than width, apex strongly constricted
(Fig. 26).
Type series.—Holotype ¢d- EARTH
School Reserve, Mpio: Pocora, Limon,
Costa Rica, August 17, 1992, R. Jones and
P. Krauter, Ex. Hampea appendiculata.
Paratypes: 2 ds, 4 2s, same data as holo-
type.
The holotype and some paratypes are de-
posited in the Collection of the Department
of Entomology. Texas A&M _ University.
Paratypes are also deposited in the Museo
Nacional de Costa Rica, San José, Costa
Rica.
Diagnosis and variation.—The long, nar-
row rostrum of the female (Fig. 3) of A.
mallyi is the most distinctive external char-
acter distinguishing this species from others
in the group. The ratio of length of rostral
apex (from antennal insertion to apex) to
width (3.6—4.2) is significantly higher for
females of A. mallyi than for the other spe-
cies. The rostra of both the males and fe-
males of this species are the least densely
hirsute of any members of the complex.
Furthermore, the rostrum of the female is
virtually devoid of distinguishable carinae
(Fig. 12). The male median lobe is uniquely
constricted at the apex (Fig. 26), readily
distinguishing this species from all others
with which it is likely to be confused.
There is little variation in the overall size
WS
of the seven specimens of the type series.
All of these specimens were reared from
flower buds of equal size under constant
conditions in the laboratory which may ac-
count for the uniformity in size. The vari-
ation noted in the color of the integument
is probably due to the teneral condition of
the reared specimens.
Host plants—Members of the type series
were reared from the flower buds of Ham-
pea appendiculata. Although populations
of H. appendiculata were examined at ele-
vations ranging from 100 to 1200 m, the
weevils were only found on trees occurring
at the lowest level in moderately disturbed
forests of the Atlantic coastal plain of Costa
Rica.
Derivation of name.—This species is
named in honor of Frederick W. Mally
(1868-1939), the first entomologist em-
ployed by the State of Texas to study the
biology and control of the boll weevil.
Anthonomus palmeri Jones and Burke,
NEw SPECIES
(Figs. 55659. 105 195 24)027,)
Body oblong-oval; length 3.42—4.92 mm
(x = 4.11, n = 30); width 1.67—2.17 mm
(x = 1.90, n = 30). Integument of body
shining, typically dark brown (some speci-
mens of type series lighter in color due to
teneral condition). Vestiture consisting of
coarse, yellowish-white pubescence; scale-
like setae somewhat broader and uniformly
distributed on thorax, occasionally more
densely concentrated on midline of prono-
tum; distributed in irregular patches on el-
ytra (Figs. 5, 6). Rostrum moderately
curved in both sexes; more strongly and
evenly curved in female (Figs. 5, 6); male
rostrum 1.7 and female 1.8 longer than
prothorax along dorsal midline. Ratio of
apical rostral length (from antennal inser-
tion to apex) to rostrum diameter (in lateral
view at level of antennal insertion) 1.6 for
male and 2.6 for female. Pubescence on
male rostrum variable in density, sometimes
sparse, extending dorsally and laterally dis-
tad of antennal insertion, that of female ex-
716
tending to antennal insertion (Figs. 9, 10).
Median and sublateral rostral carinae pro-
nounced and extending to level of antennal
insertion in both sexes. Lateral rostral
groove well defined; dorsal margin imping-
ing against eye at about middle. Eyes mod-
erately convex in dorsal view, not free be-
hind; separated in front by distance equal
to ca. ¥; width of rostrum at base. Frons
deeply foveate; covered with fairly dense
vestiture of fine, curved setae, not obscur-
ing integument. Antennae attached at about
apical ¥; of rostrum in male and at slightly
less than apical %4 in female. Funicle 7-seg-
mented; segment | slender, approximately
equal in length to next three funicular seg-
ments combined. Antennal club with basal
two segments rather loosely joined; anterior
margins of club segments bearing large
scales; apex sharply pointed; subequal in
length to preceding six funicular segments
combined. Prothorax with sides parallel to
subparallel in basal % then converging to
distinct subapical constriction; width of
apex of prothorax % width of base; integ-
ument shining, bearing deep punctures that
are smaller dorsally than laterally; areas be-
tween punctures smooth, bare. Vestiture
uniformly distributed laterally, punctures
each bearing a scale-like seta (Fig. 21),
sometimes with slight concentration of se-
tae above coxae (Fig. 17). Scutellum ca. 2X
longer than wide, sides parallel to subpar-
allel; sparsely to densely covered with fine,
appressed setae, more densely clothed at
apex. Elytra strongly convex dorsally in bas-
al 4%; distinctly wider at base than protho-
rax; humeri moderately strongly rounded;
sides subparallel to point past middle then
strongly rounded to apices; basal margin
slightly sinuate due to expanded third ely-
tral interval. Elytral pubescence arranged in
square to irregularly shaped patches on bas-
al half of elytra; coalescing into more linear
patterns towards elytral apex. Intervals
slightly convex. Ventral side of body
sparsely to densely clothed with fine to
coarse pubescence. Procoxae contiguous.
Mesocoxae separated by distance equal to
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ca. % width of coxa. Abdominal sternum 1
along midline slightly shorter than 2. Ster-
num 3 about % longer than 4. Legs mod-
erately stout (Figs. 5, 6). Profemur 3.1—
3.6X longer than broad; bearing two teeth,
the basal one being acutely pointed and
1.5-2.5xX longer than broadly triangular
distal tooth. Meso- and metafemora about
equal in size, each bearing a small, trian-
gular tooth. Protibia relatively stout, mod-
erately strongly curved in basal %; inner
margin fairly strongly sinuate. Meso- and
metatibiae not as strongly sinuate on inner
margins. Tarsal claws each bearing a long
slender tooth, the apex of which reaches
past middle of claw. Median lobe of male
slightly curved in lateral view; apex broad-
ly rounded, width equal to distance from
retracted phallotreme to apex (Fig. 27).
Type series.—Holotype d- México:
Chiapas, Tenejapa, below Paraje Yashanal,
VII-3-1990, R. W. Jones and D. Baro, ex
Hampea longipes (ground buds). Paratypes
(55) as follows: Two ds with same data as
holotype; 1 ¢, 1 2 with same label data as
holotype except collected VII-4-1990; 2 ds
with same label data as holotype except col-
lected VII-25-1990; 1 3d with same label
data as holotype except collected VI-26-
1990; 1 3 with same label data as holotype
except collected VII-14-1991 by R. W.
Jones. One 6, 2 2s-México, Paraje Yas-
hanal, Tenejapa, VIII-8-1989, R. Jones, J.
Cate, P. Krauter, ex. Hampea longipes. Two
ds, 1 ?2-Mexico-nr. Paraje Yashanal, Te-
nejapa, VII-2-1991, R. W. Jones, ex Ham-
pea longipes (ground buds). Four ds and 9
2s- México: nr. Laguna Cinco Lagos,
Mpio. La Trinitaria, Chiapas, VII-4-1990,
R. Jones and D. Baro, ex. Hampea monte-
bellensis (ground buds); 1 ¢ and 1 2 with
same data except collected VIII-2-1990; 1
2 with same label data except collected
VIII-11-1990; 3 ds and 2 2s with same
label data except collected VIII-21-1990.
Nine ds and 11 @s- México: Lagunas de
Montebello Natl. Park, VIII-9-1989, R.
Jones, J. Cate, P. Krauter, ex. Hampea mon-
tebellensis.
VOLUME 99, NUMBER 4
The holotype and some paratypes are de-
posited in the Insect Collection of the De-
partment of Entomology, Texas A&M Uni-
versity. Paratypes are also deposited in the
collections of: Instituto de Biologia, Univ-
ersidad Aut6noma de México, México, D.
FE; El Colegio de la Frontera Sur, San Cris-
tobal de las Casas, Chiapas, Mexico; Ca-
nadian Museum of Nature, Ottawa, Canada;
and Charles W. O’Brien.
Diagnosis and _ variation.—Anthonomus
palmeri is generally the smallest member of
the A. grandis complex although there is
considerable size variation in this group of
species. The small size of A. palmeri gen-
erally distinguishes it from A. townsendi
which it most closely resembles. Anthono-
mus palmeri also has a more strongly
curved rostrum with the antennae of the fe-
male attached farther from the base of the
rostrum. In addition, the profemur of A.
townsendi is more robust than that of A.
palmeri and the proximal tooth is usually
much larger than the distal tooth. The less
strongly curved median lobe with its broad-
er apex distinguishes A. palmeri from A.
grandis, A. hunteri, and A. mallyi. Further-
more, A. palmeri is separated from A. gran-
dis by having the scales on the pleural re-
gion of the prothorax uniformly distributed
as compared to the patterned arrangement
of the latter. The rostrum of the female of
A. palmeri (Fig. 5) is more hirsute and the
dorsal carinae are more distinct than in A.
mallyi or A. hunteri (Figs. 10, 12).
As is common in this species complex,
the major variation in the type series of A.
palmeri is in size. Specimens range from
3.42 to 4.92 mm in length. The diagnostic
characters are retained throughout the series
regardless of size. The greatest overall vari-
ation in size occurs between specimens of
the two host plant populations (H. monte-
bellensis and H. longipes). Specimens of A.
palmeri from H. longipes are the smallest
weevils in the A. grandis group. Specimens
from H. montebellensis are larger, though
still relatively small compared to other spe-
cies in the complex. The dorsal pronotal
WA
vitta is absent on some individuals in the
type series, most of which are from H. lon-
gipes. Scale patterns on the lateral margins
of the prothorax vary slightly in uniformity
of distribution. Some individuals have a
distinct elliptical aggregation of scales
above the procoxae as is found in A. town-
sendi; however, this aggregation is indis-
tinct in other individuals. The color of the
integument of the rostrum and prothorax
varies from brown to almost black. Al-
though there is some overlap in colors in
specimens from the two hosts, weevils
reared from H. longipes have the darkest
integument while specimens reared from H.
montebellensis are usually lighter in color.
Host plants.—The known hosts of A. pal-
meri are H. longipes in the Central High-
lands of Chiapas, and H. montebellensis in
the Lagunas de Montebello region of east-
ern Chiapas. These hosts occur above 1200
m in humid, relatively cool, montane for-
ests.
Derivation of name.—This species is
named in honor of Edward Palmer (1831-—
1911) who during his extensive natural his-
tory explorations of México was the first to
report A. grandis as a pest of cotton. After
having observed a cotton field heavily dam-
aged by the boll weevil near Monclova in
the state of Coahuila, México in 1880, he
sent specimens to entomologists in Wash-
ington, D.C. His report alerted entomolo-
gists to the threat this weevil posed for cot-
ton production.
DISCUSSION
As noted earlier, A. townsendi, A mallyi,
and A. palmeri are close allies of A. gran-
dis. In fact, they were initially considered
to be conspecific with A. grandis. However,
further analysis of their characters indicated
that they are distinct from the three recog-
nized forms of A. grandis as well as from
each other. Morphological analysis of a
large number of specimens of A. grandis
from thoughout its geographic range and
from its various hosts (Burke 1986, Burke
et al. 1986) provided a basis for critical
718
comparison of the newly discovered species
with this highly variable and widely distrib-
uted species. The most obvious and consis-
tent character separating A. grandis from
the other species of the group is the clear
demarcation of densely arranged, blunt
scale-like setae on the ventral one-half of
the pleural region of the prothorax as com-
pared to those on the dorsal one-half. This
character is present on specimens of A.
grandis from all host and geographic pop-
ulations. This, and the distinctive male me-
dian lobe, confirms that the variable popu-
lations on all hosts other than Hampea are
A. grandis. The second line of evidence that
the taxa of Anthonomus described herein
are distinct species comes from the apparent
reproductive isolation based on host and
geographic differences. Although species of
the A. grandis group are not known to oc-
cur in sympatry, large populations of A.
grandis on cultivated cotton often occur
fairly close to some of the species on Ham-
pea. For example, A. grandis was observed
to occur in large numbers in a small, iso-
lated cotton field ca. 50 kms from collection
sites of A. townsendi on H. mexicana in the
Central Depression area of Chiapas. Given
the proven flight capabilities and abundance
of A. grandis, the probability of eventual
sympatry occurring between this species
and A. townsendi would appear to be high,
especially since H. latifolia is known to
serve as host for both species. However, no
sympatry or evidence that the two species
interbreed have been found.
The discovery of three undescribed spe-
cies of Anthonomus on four of the nine spe-
cies of Hampea examined in the field re-
veals some interesting information about
the association of species of the complex
with their Hampea hosts. The restriction of
A. townsendi, A. mallyi, A. palmeri, and the
previously described A. hunteri to Hampea
as hosts, and the fact that species of this
plant genus also serve as occasional hosts
of A. grandis, strongly supports the conclu-
sion of Burke et al. (1986) that Hampea is
the ancestral host genus of the entire group.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Since Hampea and members of the A. gran-
dis group have apparently had a long as-
sociation, the possibility is great that other
undescribed species of related weevils re-
main to be discovered on the remaining
eight species of Hampea not examined dur-
ing this and previous studies. However, the
fact that weevils of this group were not
found on five species of Hampea examined
in the field indicates that ecological, mor-
phological, historical, and/or chemical fac-
tors may be involved in preventing infes-
tations. Further field and taxonomic studies
of both the A. grandis species complex and
the genus Hampea may elucidate the fac-
tors involved and nature of the evolutionary
processes in this insect/plant association.
There is considerable urgency in conduct-
ing such studies since some species of
Hampea, and possibly their associated wee-
vils, may have recently been extirpated and
others are clearly threatened.
ACKNOWLEDGMENTS
The authors thank Deborah Baro of Que-
rétaro, México, James Cate of Lawrence-
burg, Indiana, and Pete Krauter of Texas
A&M University for help in making field
collections and for assistance with various
other aspects of this research. Paul Fryxell,
University of Texas, helped immensely in
the process of locating and identifying the
species of Hampea studied here. Wayne
Clark, Auburn University, provided useful
comments on the manuscript. Photomicro-
graphs were made at the Electron Micros-
copy Center at Texas A&M University.
Collections in Costa Rica were made pos-
sible with the help of the Organization of
Tropical Studies, San José, Costa Rica in
obtaining a collecting permit and with co-
operation from the EARTH School in Po-
céro, Costa Rica. This study was conducted
with funds provided by the Texas Agricul-
tural Experiment Station, with support from
USDA Science and Education Grant
85-CRCR-1-1688, and the USDA Foreign
Agricultural Service, International Cooper-
VOLUME 99, NUMBER 4
ation and Development Grant FG-MX-102,
Project MX-ARS-3.
LITERATURE CITED
Bodegas, V., P. R., G. R. Flores, and M. E. de Coss
Flores. 1977. Aspectos de interés sobre las hos-
pederas alternantes del picudo del algodonero A.
grandis y avances en la investigacion respectiva
en el Soconuso, Chiapas, Mexico. Ecoldogicas del
Sureste, Boletin 3, 14 pp.
Burke, H. R. 1986. Situacao taxondmica do bicudo
do algodoeiro no Brasil e em outras areas da
America do norte e do sul, pp. 89-134. In O Bi-
cudo do Algodoeiro. Empresa Brasileira de Pes-
quisa Agropecudria, EMBRAPA, Departamento
de Difusao de Technologia, Brasilia, D.F, 314 pp.
Burke, H. R., and J. R. Cate. 1979. A new species of
Mexican Anthonomus related to the boll weevil
(Coleoptera: Curculionidae). Annals of the Ento-
mological Society of America. 72: 189-192.
Burke, H. R., W. E. Clark, J. R. Cate, and P. A. Fryxell.
1986. Origin and dispersal of the boll weevil.
Bulletin of the Entomological Society of America.
32: 228-238.
Cate, J. R., PB C. Krauter, and K. E. Godfrey. 1990.
Pests of cotton, pp. 17-19. In Habeck, D. H., FE
D. Bennett, and J. H. Frank, eds. Classical Bio-
logical Control in the Southern United States.
Southern Cooperative Series Bulletin No. 355.
Clark, W. E. 1988. Revision of the weevil genus Lon-
cophorus Chevrolat (Coleoptera: Curculionidae,
WM
Anthonominae). Quaestiones Entomologicae 24:
465-518.
1995. A new species of Loncophorus from
Mexico (Coleoptera:Curculionidae, Anthonomini)
associated with Mortoniodendron (Malvales:Tili-
aceae). Anales del Instituto de Biologia, Univer-
sidad Nacional Aut6noma de México, Serie Zoo-
ld6gica. 66: 107-112.
Clark, W. E., and H. R. Burke. 1986. Phylogeny of
the species of the Anthonomus subgenus Antho-
nomorphus Dietz, with discussion of relationships
with Anthonomus grandis Boheman (Coleoptera:
Curculionidae). Journal of the Kansas Entomolog-
ical Society. 59: 508-516.
Cross, W. H., M. J. Lukefahr, P. A. Fryxell, and H. R.
Burke. 1975. Host plants of the boll weevil. En-
vironmental Entomology. 4: 19-26.
Fryxell, P A. 1969. The genus Hampea (Malvaceae).
Brittonia 21: 359-396.
1979. The Natural History of the Cotton
Tribe. Texas A&M University Press, College Sta-
tion, Texas, 245 pp.
Fryxell, P. A., and M. J. Lukefahr. 1967. Hampea
Schlecht: possible primary host of the boll weevil.
Science 155: 1568-1569.
Stansly, P. A. 1985. Life history and ecology of the
boll weevil on a native host plant in southeastern
Mexico, Ph.D. Dissertation, Texas A&M Univer-
sity, College Station, Texas.
Townsend, C. H. T. 1895. Report of the Mexican boll
weevil in Texas (Anthonomus grandis Boh.). In-
sect Life 7: 259-309.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 720-726
A NEW SUBGENUS AND SPECIES OF THE GENUS BRACHYUNGUIS DAS
(HEMIPTERA: APHIDIDAE) FROM ARGENTINA
M. PILAR MIER DURANTE, JAIME ORTEGO, AND JUAN M. NIETO NAFRIA
(MPMD, JMNN) Departamento de Biologia Animal, Universidad de Leon, E-24071
Leén, Spain; (JO) AER INTA Malargiie, Saturnino Torres 862, 5613 Malargiie (Mza.),
Argentina.
Abstract.—Brachyunguis (Andinaphis) paradoxus, new Argentinean subgenus and
species are described. The new subgenus and new species are characterized by the absence
of marginal papillae, processus terminalis of the antennal segment VI longer than the base
of segment, tarsal formula 3.3.3 and its host-plant, Senecio (Asteraceae). These characters
are very atypical among Brachyunguis and even Aphidini. Apterous and alate viviparous
females and oviparous females are described. The subgenus and species are keyed in two
modified previous keys.
Key Words: aphid, Brachyunguis, Andinaphis, new subgenus, new species, Senecio, Ar-
gentina
Resumen.—‘‘Nuevos subgénero y especie del género Brachyunguis Das (Hemiptera
Aphididae) de Argentina’. Se describen las hembras viviparas apteras y aladas y las
oviparas de Brachyunguis (Andinaphis) paradoxus n. subgen., n. sp., a partir de ma-
terial recogido sobre Senecio subumbellatus (Asteraceae) en la provincia de Mendoza
(Argentina). La completa ausencia de papilas marginales en el prot6rax y en el abdomen
(caracter atipico en Brachyunguis y muy infrecuente en Aphidini), la longitud del fila-
mento terminal del VI artejo antenal: mas largo que la base del mismo, la formula tarsal:
3.3.3, y la planta hospedadora, caracterizan tanto el subgénero como la especie. Ambos
nuevos taxones se incluyen en conocidas claves anteriores, que se modifican convenien-
temente.
Palabras clave: pulgén, Brachyunguis, Andinaphis, subgénero nuevo, especie nueva,
Senecio, Argentina
Two species of Brachyunguis Das, 1918 In North America (Remaudiére and Hal-
are known in South America: B. (Xerophi-
laphis) blanchardi Remaudiére and Baha-
mondes, 1987, and B. (B.) bahamondesi
Remaudiére and Halbert, 1996. Both were
described recently from Mendoza province,
Argentina, and live on Chenopodiaceae, on
Allenrolfea vaginata and on Atriplex lampa
plus Chenopodium sooanum, respectively.
bert 1996) three other species of the genus
are known, all of them on Chenopodiaceae:
B. (B.) bonnevillensis (Knowlton, 1928) on
Atriplex and Sarcobatus, B. (X.) tetrapter-
alis (Cockerell, 1902) on Atriplex and Suae-
da, and B (B.) bishopi Remaudiere and Hal-
bert, 1996 on Sarcobatus.
The plant hosts of the Brachyunguis spe-
VOLUME 99, NUMBER 4
cies of the Old World are Chenopodiaceae
(mainly), Tamaricaceae, Asclepiadaceae,
Zygophyllaceae and Sapindaceae, never
Asteraceae.
The genus Brachyunguis is included in
the subtribe Aphidina. Marginal papillae
are present normally on the prothorax and
abdominal segments I and VII, and fre-
quently also on abdominal segments II to
VI in this subtribe. However, there are some
exceptions (Eastop 1979): Cryptosiphum
Buckton, 1879 (on Artemisia, Asteraceae,
palearctic), Indiaphis Basu, 1969 (on Rho-
dodendron, Ericaceae, oriental) but this ge-
nus may be included in Macrosiphini (Re-
maudiére and Remaudiére 1997), Mistura-
phis Robinson, 1967 (on Artemisia, Aster-
aceae, nearctic), Polytrichaphis Miyazaki,
1971 (on TIllicium, Wliciaceae, palearctic
from Japan, also included in Macrosiphini
by Remaudiére and Remaudiere 1997) and
Sanbornia Baker, 1920 (on Juniperus, Cu-
pressaceae, nearctic).
Brachyunguis is divided in two subgen-
era Brachyunguis s. str. (present in Asia,
Europe, North [Mediterranean] Africa and
America, and Xerophilaphis Nevsky, 1928
(Asia and America). Also from Mendoza
province (Argentina), the second author
found some aphids which can be ascribed
to the genus Brachyunguis. However, they
belong to a new subgenus, Andinaphis,
characterized by the absence of marginal
papillae (an atypical characteristic in Aphid-
ini) and three atypical characteristics among
Brachyunguis: a) processus terminalis of
the antennal segment VI longer than the
base of segment, b) tarsal formula 3.3.3,
and c) unusual host-plant, Senecio (Aster-
aceae).
Abbreviations used in the text are as fol-
lows: ant.III, IV, V = antennal segments III,
IV, V; ant. VIb = base of antennal segment
VI; ant.VIpt = processus terminalis of an-
tennal segment VI; b.d.IIJ = basal diameter
of antennal segment III; BL = body length;
h.t.II = second segment of hind tarsus; u.r.s.
= ultimate rostral segment.
721
Andinaphis Mier Durante,
Ortego, and Nieto Nafria,
NEW SUBGENUS
Type species.—Brachyunguis (Andina-
phis) paradoxus nN. sp.
Etymology.—The name Andinaphis is
taken from the Neo-Latin words ‘“‘andinus”’
and “‘aphis’’, Andean and aphid, respec-
tively. The gender is feminine as Aphis.
Description.—Light green or yellow
when alive, poorly pigmented, without seg-
mental sclerites on abdomen. Frontal profile
convex, antennae shorter than body length,
processus terminalis of antennal segment
VI longer than 1.3 times base of this seg-
ment, secondary rhinaria normally present
on antennal segment III of apterous vivip-
arous females and on antennal segments III
and IV of alatae, also sometimes present on
III on oviparous females, tarsal formula
3.3.3 with exceptions (2 setae on front and
middle legs), siphunculi truncated-cone
shape and shorter and darker than cauda,
which is more or less long triangular or fin-
gerlike with 8 setae at most, without mar-
ginal papillae on prothorax and abdominal
segments, including I and VII. Monecious
holocyclic on Senecio (Asteraceae), on ae-
rial parts of the plants.
Discussion.—Andinaphis can be distin-
guished from other subgenera of Brachyun-
guis (Brachyunguis s. str. and Xerophila-
phis) by: 1, the total absence of marginal
papillae (marginal papillae are present on
prothorax and abdominal segments I and
VII in both other subgenera); 2, the proces-
sus terminalis of VI antennal segment is
1.3—1.9 times as longer as base of the seg-
ment (this ratio is normally smaller in other
subgenera); 3, the tarsal formula is
3(2).3(2).3, in the other subgenera it is 2.2.2
or 2(3).2(3).2; and 4, Asteraceae, not Che-
nopodiaceae, is the host-plant.
Andinaphis can not be confused with
Protaphis (subgenus of Aphis) which also
lives on Asteraceae and has small processus
terminalis (0.9—1.5X base), because of five
important characters (see the respective
V22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
characters on Andinaphis in the descrip-
tion). Protaphis has mainly tarsal formula
3.3.2, is dark brown to black alive with
black or very dark cauda, has some seg-
mental sclerites on abdomen of apterous fe-
males, has 7—16 caudal setae, and often
lives on the subterranean parts of the plants.
The subgenus can be identified by mod-
ifying Eastop’s (1979) key as follows.
1—4. [without modification]
3D) Lateral tubercles [papillae] absent from both
first and seventh and usually also the other
abdominal segments. Processus terminalis
1.3—5.5 times as long as the base of the sixth
antennallscorneniiquesn t-te ite enneee er 6
- Lateral tubercles present on the first and
usually also on the seventh abdominal seg-
IMMETMUS Fs Sho cc wetege pose OE ec casa okes oe ics 8
6. Triommatidion inconspicuous. Siphunculi
wider than and about 2% times as long as
the cauda. On Rhododendron (Ericaceae),
India [in Macrosiphini, by Remaudiére and
RemardierceliO 97) ees aeneae een Indiaphis
- Triommatidion evident at hind margin of eye
OOP On tos ee 6b
6b. All first tarsal segments bearing only 2 hairs.
Processus terminalis 3.0—5.5 times as long
aSutnelbaserom antennal V/s sei eiel ene ae 7
- Normally first tarsal segments bearing 3
hairs, sometimes some tarsus of the front
and medium legs with 2 hairs. Processus ter-
minalis 1.3—1.9 times as long as the base of
antennal VI. On Senecio (Asteraceae), South
/NANETICEY oo oo ceo oe Brachyunguis (Andinaphis)
U- (and the following disjunctives) [without varia-
tion]
Brachyunguis (Andinaphis) paradoxus
Mier Durante, Ortego, and Nieto
Nafria, NEW SPECIES
(Bigsgulee2)
Apterous viviparous female (n = 192; 31
measured) (Figs. 1, 2).—Body 1.11 to 2
mm long. Light green when alive, excep-
tionally yellow greenish, with white waxy
powder; pale specimens with only tarsi,
knees, apex of siphunculi and ultimate ros-
tral segment light brown and more pig-
mented specimens with a part of femora,
knees, apex of tibiae, tarsi, ultimate rostral
segment, siphunculi and anal plate brown
and cauda light brown. Prepared specimens
light yellow in general with head, distal ros-
tral segments, legs (apex of tibiae and tarsi
browner), siphunculi, cauda and genital and
anal plates more or less light brown and
intersegmental (sometimes inconspicuous)
and stigmatic sclerites smoky.
Cuticle more or less smooth. Setae (table
1) pale and acute in general. Without pa-
pillae on prothorax and abdominal seg-
ments I and VII. Frontal profile convex. Se-
tae on vertex usually shorter than b.d.III
and longer than antennal setae. Clypeus
more or less as wide as interantennal space
but never exceeding front. Antenna 6 seg-
mented, (0.58)0.7 to 1.03 mm long, 0.45 to
0.67 times body length (BL); ant.III with
(0)1—6(8) widely irregular and flanged sec-
ondary rhinaria; antennal segment lengths
Gan mm): ant.JIII = (0.13)0.16 to 0.29;
ant.IV = (0.09)0.11 to 0.19; ant.V = 0.09
to 0.18; ant. VIb = 0.07 to 0.10; ant. VIpt =
(0.10)0.12 to 0.17; ant.III 1.20 to 2.13 times
longer than ant.VIpt, which is 1.33 to 1.88
times longer than ant.VIb; antennal setae
few (2 to 6 on ant.III and 1 to 3 on ant. VI.b)
and short. Rostrum (0.38 to 0.47 mm long)
reaching third coxa; BL 2.54 to 4.65 times
length of rostrum; u.r.s pointed, with sides
straight or slightly concave, 0.08 to 0.11
mm long, 1.54 to 2.11 times as long as its
basal width, 0.70 to 0.80(0.95) longer than
h.t.II, 0.90 to 1.12 times ant.VIb., with 2
accessory lateral setae. Posterior seta on
trochanter of hind leg shorter than the basal
diameter of femur; dorsal and ventral setae
on femora few (2 to 6 ventral ones) and
short; hind tibia 0.34 to 0.55 times BL; first
tarsal segments with 3.3.3 setae, exception-
ally 2 setae on meso- and meta-thoracic
legs; h.t.II 0.10 to 0.14 mm long, 0.15 to
0.25 times hind tibia. Abdominal setae
short; presiphuncular abdominal segments
with 2 marginal setae each side and nor-
mally with 2 spinal setae; abdominal seg-
ment VIII with 2 setae, exceptionally 4,
longer than anterior ones. Siphunculi trun-
cated-cone shape, 0.05 to 0.10 mm long,
0.95 to 1.54(1.73) times its basal width and
VOLUME 99, NUMBER 4 723
SB= Es 0.5mm.
Fig. 1. Brachyunguis (Andinaphis) paradoxus, habitus. Illustration by M. Enrique Ortega Lorenzo (Leon,
Spain).
724 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Brachyunguis (Andinaphis) paradoxus. A, Ill antennal segment. B, VI antennal segment. C, Ultimate
rostral segment. D, Hind tarsus. E, Siphunculus. K Cauda. Illustrations by M. Enrique Ortega Lorenzo (Leon,
Spain).
Table 1. Measurements of setae of Brachyunguis (Andinaphis) paradoxus. Abbreviations: D = diameter of
trochanter-femoral joint; abd. seg. = abdominal segment; other abbreviations as in text.
ap. viv. fem al. viv. fem. ovip. fem.
ant. III p.m long 7-13 10-13 7-10
b.d. Ill times 0.40.8 0.6—1.0 0.4—-0.7
Vertex wm long 10-18 10-20 11-14
b.d. II times 0.6—1.3 (0.8)1.0-1.6 0.7-1.0
Hind trochanter wm long 20-30 22-30 22-33
D times 0.5-0.8 0.5—0.7 0.5-0.8
Dorsal hind femur ym long 5-18 12—23 7-18
b.d. II times 0.4-1.3 (0.7)1.0-1.8 0.4—-1.2
Ventral hind femur ym long 12-25 17-28 18-24
b.d. III times 0.7-1.8 1.3—2.0 1.0-1.6
abd. seg. II: spinal wm long 10-14 10-18 11-15
abd. seg. III: marginal ym long 10-18 12-20 8-15
b.d. II times 0.6-1.1 0.6—1.5 (0.4)0.6—1.0
abd. seg. VIII wm long 12-25 15-28 15-23
b.d. ILI times 0.9-1.6 1.0—2.0 1.0-1.7
VOLUME 99, NUMBER 4
0.33 to 0.52(0.62) times the cauda; subgen-
ital plate with 2(4) anterior, 0(2) discal and
5—10 posterior setae; cauda fingerlike,
(0.10)0.13 to 0.20 mm long, 1.40—1.78 lon-
ger than its basal width, with 5 to 7 lateral
and exceptionally 1 dorsal setae.
Alate viviparous female (n = 38; 24
measured).—Body (1.12)1.52—1.89 mm
long. Alive and mounted similar to the ap-
terae, but darker on head, antenna (some
times the basal art of ant.II]), thorax and
legs. Metric and meristic characters very
similar to aptera, but with following differ-
ences. Antenna = 0.80—1.06 mm, 0.51-—
0.68 times BL; secondary rhinaria: 4—9 on
ant.III and O(most frequent)—2 on ant.IV;
antennal segment lengths (in mm): ant.III
= 0.21-0.31, ant.IV = 0.17—0.22, ant.V =
0.12—0.24, ant.VIb = 0.8—0.12; ant.VIpt =
0.13-0.18; ant.HI 1.33-1.94 times
ant.VIpt.; u.rs. 0.82—-1.05 times ant.VIb;
u.r.s. 0.64—0.85 times h.t.II; siphunculus
1.16—2 times its basal width; subgenital
plate with 2—5 anterior, O—2 discal and 6—9
posterior setae; cauda |.17—1.70 times its
basal width. Setae (Table 1) similar to those
in apterae.
Oviparous female (n = 35; 14 mea-
sured).—Body 1.4 to 1.8 mm long. Very
similar to viviparous female, with metatho-
racic tibiae enlarged with (40)60 to 110
scent plates. But some characters slightly
different: hind tibia 0.28—0.40 times BL;
antenna (0.64—0.95 mm long) 0.41—0.67
times BL; ant.III and ant.[1V secondary rhi-
naria 0—1(4) and O, respectively; ant.III
1.06 to 1.74 times ant.VIpt; u.rs. 0.67—0.9
times h.t.II; 4 to 10 setae on abdominal ter-
gite VIII; subgenital plate with 11—28 an-
terior and discal (mixed) and 10—16 poste-
rior setae; cauda 1.05—1.67 longer than its
basal width with 4-8 lateral setae. Setae
(Table 1) as in apterae ones.
Eggs measured in the females: 0.7 X 0.4
mm.
Male.—Unknown.
Type material.—Holotype: apterous vi-
viparous female (measured specimen num-
ber 1) collected on Senecio subumbellatus
725
Phil. at Malargiie “Arroyo El Rezago”
(Mendoza province, Argentine), 8-III-96, J.
Ortego leg., in collection Universidad de
Leén (Departamento de Biologia Animal).
Paratypes: apterous, alate viviparous and
Oviparous females found (J. Ortego leg.) on
the same host-plant at the same locality,
8-I1I-96 and 7-IV-96 and on the same host-
plant at Malargiie ““‘Refugio del Club An-
dino”’, 15-V-94, deposited in the authors’
collections and in The Natural History Mu-
seum, London, Muséum _ National
d’Histoire Naturelle, Paris, and S. Halbert
collection, Gainesville, Florida (USA).
Etymology.—The specific name is taken
of classic Greek: Tapadoéoo, paradoxical,
surprising, because the peculiar characters
of the species (see discussion of the new
subgenus).
Biology and distribution.—Brachyunguis
(Andinaphis) paradoxus is monoecious
holocyclic on Senecio subumbellatus and
perhaps on other Andean Senecio spp. It
forms small and dense colonies on the ae-
rial part of the plant, mainly on the floral
stems. It is very possible that this new spe-
cies is restricted in distribution to the Andes
Range in Argentinian provinces of Men-
doza (Malargtie: “‘Arroyo El Rezago’’,
1905 m, “Refugio del Club Andino”’, 2225
m and “El Carrizalito”’, 2025 m) and Neu-
quen plus adjacent localities of Chili, as is
its plant-host.
Discussion.—Brachyunguis (Andinaphis)
paradoxus is the only known species of the
subgenus. The key established by Remau-
diére and Halbert (1996) to the American
species of Brachyunguis can be modified to
include the new species as following form:
1. [without variation]; subgenus Xerophilaphis
— [without variation]
2. [without variation, to B. (X.) tetrapteralis
and B. (X.) blanchardi)
2b. Marginal papillae absent on prothorax and
abdomen. Ant.VIpt 1.33 times ant.VIb at
least and normally more than 1.4; urs.
shorter than h.t.Ifl. On Senecio. Argentina;
subgenus Andinaphis: ..... B. (A.) paradoxus
— Marginal papillae present on prothorax and
abdominal segments I and VII. Ant.VIpt
1.30 times ant.VIb normally, if 1.4 times,
u.r.s. longer than h.t.II. Not on Senecio;
BrAGHYUNSUISES Site ee au eee eee 3
[without variation, to B. (B.) bishopi and B.
(B.) bonnevillensis]
3-6.
ACKNOWLEDGMENTS
The authors express their gratitude to
Prof. Georges Remaudiére (Muséum Na-
tional d’Histoire Naturelle) for his com-
ments and news from his Catalogue then in
press, Dr. Susan Halbert (Florida Depart-
ment of Agriculture and Consumer Ser-
vices) for her comments and for reviewing
the manuscript, and Prof. Gaspar Morocho
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
for his assistance in the etymology of new
taxa names.
LITERATURE CITED
Eastop., V. EF 1979. Key to the genera of the subtribe
Aphidina (Homoptera). Systematic Entomology 4:
379-388.
Remaudiére, G. and L. Bahamondes. 1987. Un
Brachyunguis (Xerophyllaphis) nouveau d’ Argentine
(Hom. Aphididae). Revue frangaise d’Entomologie
(Nouvelle Serie) 9(4): 179-185.
Remaudiére, G. and M. Remaudiére 1997. Catalogue
des Aphididae du Monde/Catalogue of the world’s
Aphididae. Homoptera Aphidoidea. Institut National
de la Recherche Agronomique. Paris. 475 pp.
Remaudiére, G. and S. Hallbert 1996. American spe-
cies of the genus Brachyunguis Das (Homoptera:
Aphididae), including two new species. Proceed-
ings of the Entomological Society of Washington
98(2): 249-255.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 727-743
ANATOMY OF THE REPRODUCTIVE SYSTEM IN SIX ANASTREPHA
SPECIES AND COMMENTS REGARDING THEIR TERMINOLOGY IN
TEPHRITIDAE (DIPTERA)
IMELDA MARTINEZ M. AND VICENTE HERNANDEZ-ORTIZ
(IMM) Departamento de Ecologia y Comportamiento Animal, Instituto de Ecologia y
Comportamiento Animal, Instituto de Ecologia, A.C., Apartado Postal 63, Xalapa, Vera-
cruz 91000, Mexico; (VH) Departamento de Biosistematica de Insectos, Instituto de Ecol-
ogia, A.C., Apartado Postal 63, Xalapa, Veracruz 91000, Mexico
Abstract.—A comparative study of the anatomy of the reproductive system of mature
males and females of the following species was performed: A. serpentina (Wiedemann),
A striata Schiner, A cordata Aldrich, A. ludens (Loew), A. obliqua (Macquart) and A.
fraterculus (Wiedemann). The study material came from diverse regions of the states of
Veracruz and Chiapas, Mexico. In females, several important differences among species
were observed: in the number of ovarioles, morphology of the ventral receptacle, signum,
spermathecae and aculeus. In males, differences were apparent in the accessory glands,
ejaculatory apodeme of the sperm pump, distiphallus and surstyli. For the first time, the
real location of the seminal vesicles is shown and the aedeagal gland is described. A
comparative discussion ensues regarding terminology and the location of various struc-
tures in both sexes.
Key Words:
The genus Anastrepha includes 185 spe-
cies occurring only in the Western Hemi-
sphere. Despite the great diversity and wide
distribution of this genus, most studies have
concentrated exclusively on a few species
of economic importance (Hernandez-Ortiz
and Aluja 1993).
Williamson’s (1989) review and synthe-
sis of studies of the descriptive anatomy of
the reproductive system, oogenesis and
spermatogenesis was limited to only a few
fruit fly species. Studies of females of the
following species have been published:
Rhagoletis pomonella (Walsh) (Dean 1935),
Ceratitis capitata (Wiedemann) (Hanna
1938, Valdéz Carrasco and Prado Beltran
1990), Xanthaciura unipuncta Malloch
(Souza Lopes 1939), Bactrocera tryoni
(Froggatt) (formerly Strumeta) (Drew
Anastrepha, reproductive system, females, males, anatomy
1969), Bactrocera oleae Gmelin (formerly
Dacus) (Solinas and Nuzzaci 1984), as well
as Rhagoletis nova (Schiner) and R. con-
versa (Brethes) (Flores et al. 1987).
For Anastrepha there is only general in-
formation regarding the anatomy of the re-
productive system in females for A. /udens
(Loew) (Dampf 1933, Servin-Villegas and
Jiménez-Jiménez 1995), for A. suspensa
(Loew) (Dodson 1978) and for A. serpen-
tina (Wiedemann) (Martinez et al. 1995).
With respect to males, the information
presently available consists of studies in C.
capitata (Hanna 1938, Anwar et al., 1971),
X. unipuncta (Souza Lopes 1939) and B.
tryoni (Drew 1969). The studies of Anas-
trepha include species such as the follow-
ing: A. suspensa (Dodson 1978), A. serpen-
tina (Martinez et al. 1995), A. ludens (Ser-
728
vin-Villegas and Jiménez-Jiménez 1995)
and several unidentified Anastrepha species
(Bressan 1995).
A review of the aforementioned literature
shows that much of the terminology used
for Tephritidae by these authors differs sub-
stantially, regarding names and locations of
certain structures.
In this paper we present a comparative
study of the anatomy of the reproductive
system for males and females of the follow-
ing species: A. serpentina (Wiedemann), A.
striata Schiner, A. cordata Aldrich, A. lu-
dens (Loew), A. obliqua (Macquart) and A.
fraterculus (Wiedemann). Furthermore, ter-
minology and the location of certain struc-
tures such as the seminal vesicles and the
aedeagal gland in males, and the bursa cop-
ulatrix and the ventral receptacle in females
are discussed.
MATERIAL AND METHODS
Males and females of six Anastrepha
species were obtained from larvae infesting
their natural host plants from the following
regions: A. serpentina from Region del So-
conusco, Chiapas reared in Pouteria sapota
(Sapotaceae); A. striata and A. fraterculus
from the “‘Estacién de Biologia Tropical
Los Tuxtlas’’, Veracruz reared in Psidium
guajava (Myrtaceae); A. cordata from the
“Estacion de Biologia Tropical Los Tux-
tlas’’, reared in Tabernaemontana alba
(Apocynaceae); A. /udens from Martinez de
la Torre, Veracruz, reared in Citrus sinensis
(Rutaceae); and A. obliqua from Apazapan,
Veracruz, reared in Spondias purpurea (An-
acardiaceae).
The emergence and development of
adults took place under laboratory condi-
tions; until the gonads reached full maturi-
ty, hydrolized protein (ICN-Biomedial Inc.)
was provided as a food source. Approxi-
mately 50 individuals of each species and
sex were analyzed (except A. cordata, for
which only 7 specimens were observed).
Dissection of the individuals was _per-
formed under a stereomicroscope. Each
specimen was placed in a petri dish in
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Ringer’s solution. The reproductive systems
were drawn while in this solution using a
camera lucida. Specimens were fixed on
slides in Carnoy’s solution, then dehydrat-
ed, cleared and stained using the Feulgen
light green technique. Finally, they were
mounted whole in Canada Balsam (Gabe
1968).
In order to observe particular morpholog-
ical structures such as the cuticular intima
of the vaginal duct, ventral receptacle, sper-
mathecae, and the aculeus in females, as
well as the vas deferens, accessory glands
and sperm pump in males, some reproduc-
tive systems were dissected in NaOH so-
lution (4%) and stained with chlorazol
black E following the technique described
by Carayon (1969). Later they were dehy-
drated, cleared and mounted in Canada Bal-
sam.
The terminalia of both sexes were pre-
pared by cutting open the entire abdomen
in NaOH (10%) solution which was boiled
for a few minutes in order to remove all
extraneous materials; the terminalia were
later analyzed with a compound micro-
scope. Microphotographs were made with a
Zeiss photomicroscope.
RESULTS
Female Reproductive System
The general terminology employed here
to describe females is based on the works
of McAlpine (1981) and Norrbom and Kim
(1988), and for particular internal structures
we following Dean (1935). There is consid-
erable terminological confusion among lat-
er studies that used different anatomical
terms for the same structure or used incor-
rect terminology (Table 1).
The female Anastrepha reproductive sys-
tem is made up of the following structures:
two ovaries, two lateral oviducts, the com-
mon oviduct, two accessory glands, three
spermathecae with their respective ducts,
vagina and aculeus (Fig. 1A).
Even in mature females, the ovary is
covered by a thin wall. The ovarioles (Fig.
VOLUME 99, NUMBER 4 729
Table 1. Comparison of some terminology used for the female reproductive system in Tephritidae.
Present Study Dampf (1933) Dean (1935) Hanna (1938) Drew (1969) Solinas & Williamson
Nuzzaci (1984) (1989)
Ovary ovary ovary ovary ovary ovary ovary
Lateral oviduct not indicated lateral oviduct oviduct lateral oviduct lateral oviduct lateral oviduct
Common ovi- not indicated median ovi- common ovi- not indicated common ovi- common ovi-
duct duct duct duct duct
Accessory accessory accessory colleterial colleterial accessory accessory
glands glands glands glands glands glands glands
Spermathecae spermathecae spermathecae spermathecae spermathecae receptaculum
seminis spermathecae
Bursa copulatrix not indicated bursa copu- vagina vagina anterior vagi- bursa copu-
latrix na latrix
Ventral recepta- not indicated ventral recep- vagina morula gland fertilization ventral recep-
cle tacle
Vaginal duct oviduct vagina
Cloacal opening end oviduct cloaca
vaginal duct
genital open-
ing
vaginal duct
gonopore
chamber
posterior va-
gina
not indicated
tacle
vaginal duct
vulva
1B) are of the polytrophic type and have a
terminal filament, germarium, vitellarium
and calyx. The germarium contains the
odgonial cells, nutritive cells and prefollic-
ular cells. The follicles found in the vitel-
larium show varying degrees of maturity;
each one is formed by 16 cells of germinal
origin corresponding to one oocyte and 15
nutritive cells which are contained in the
cavity defined by the follicular epithelium.
The calyx, which opens into its respective
lateral oviduct, is a prolongation of the
vitellarium. The lateral oviducts are short
and their walls very thin, both open into the
common oviduct, which is slightly longer
and reaches the anterior part of the vagina.
The vagina has two regions with differ-
ent characteristics. The anterior part is mod-
ified to form a differentiated region of dis-
tinctive structure called the bursa copu-
latrix. The posterior part or vaginal duct is
a long tubular duct which folds up before
penetrating the aculeus. Parallel to the rec-
tum, it flows into the cloaca.
The bursa copulatrix can be observed
externally due to the greater thickness of its
wall and because both the spermathecal and
accessory gland ducts open into its dorsal
side. The thickness of the bursa copulatrix
is partly due to the presence of the ventral
receptacle, a flatttened, sac-like structure
protruding cephalad from its ventral side.
The ventral receptacle has thick, highly or-
namented cuticular walls.
The vaginal duct is long and tubular and
it has a number of longitudinal pleats. At
the end of its basal third, positioned dor-
sally, are two cuticular pieces called the sig-
num.
Both accessory glands are oval in shape,
voluminous and opaque. Each gland has a
bulky base and extends into a thin, long
duct which opens into the anterior part of
the vagina.
The spermathecae are each made up of a
sclerotized capsule surrounded by a secre-
tory epithelium connected by a very long
and thin spermathecal duct. All ducts open
into the dorsal side of the bursa copulatrix.
The aculeus is a highly sclerotized struc-
ture; its morphology and length differ
among species of Anastrepha, making it ex-
tremely important for the taxonomy of the
genus.
In the studied Anastrepha species, the fe-
male reproductive system varies mainly in
the number of ovarioles and in the morpho-
logical characteristics of the ventral recep-
tacle, as well as in the signum, spermathe-
cae and aculeus.
730
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
+eva
Fig. 1. Reproductive system of the female of Anastrepha serpentina. A, General morphology of the repro-
ductive system. B, Microphotograph of the apical end of some ovarioles. Abbreviations: ag = accessory gland;
be = bursa copulatrix; bo = basal mature oocyte; co
= common oviduct; eva = entrance of the vagina to
aculeus; ge = germarium; lo = lateral oviduct; of = ovarian follicles in the vitellarium; ov = ovary (with some
ovarioles figured); si = signum; sp = spermatheca; vgd = vaginal duct; vr = ventral receptacle.
The number of ovarioles present in the
Six species varied between 20 and 33 ovar-
ioles per ovary. The highest average was
observed in A. obliqua (30-33), A. serpen-
tina (25-30) and A. cordata (25—28); the
lowest averages were in A. fraterculus (20—
24), A. striata (20—25) and A. ludens (22—
25) (fable).
The ventral receptacle is usually elongat-
ed with numerous lobular papillae on its
surface (Fig. 2A,B,D,E,F) except in A. cor-
data, in which it is almost spherical, shorter
and almost completely without papillae
(Fig: 2C; Table: 2).
The signum is formed by two sclerotized
plates which are long and sclerotized in A.
serpentina and A. striata (Fig. 2A’, B’),
short and less sclerotized in A. ludens (Fig.
2D’,E’), very short and more or less scler-
otized in A. fraterculus (Fig. 2F’) and in-
conspicuous in A. cordata (Fig. 2C’).
The glandular units of the spermathecal
secretory epithelium differ among species
(Fig. 3). The spermathecal capsules are
spherical in shape and usually covered by
numerous sclerotized spiculae. The sper-
mathecae of A. Judens tend to be longer
than those of other species. In A. cordata,
in contrast, the spermathecae are rounded
and the spiculae are lacking. In A. frater-
VOLUME 99, NUMBER 4
Table 2. Some comparative characteristics in fe-
males of the Anastrepha species examined during this
study.
Number Ventral
Species Ovarioles Receptacle Signum
A. serpentina 25-30 elongated 2 plates
A. striata 20-25 elongated 2 plates
A. cordata 25-28 short inconspicuous
A. ludens 22-25 elongated 2 plates
A. obliqua 30-33 elongated 2 plates
A. fraterculus 20-24 elongated 2 plates
culus the spermathecae are not consistently
rounded, as they are in A.obliqua (Fig. 4
A-F).
The aculeus is usually sclerotized and
dorsoventrally compressed. In A. serpentina
it measures from 2.9—3.3 mm long; the tip
(Fig. 4A’) has tiny teeth which occupy the
apical third. In A. striata, the aculeus is no-
tably robust, measuring 2.1—2.3 mm long;
its tip (Fig.4B’) is broad with a marked con-
striction after the cloacal opening. The acu-
leus of A. cordata is very thin (less than
0.09 mm wide) and measures 4.7—5.5 mm
long; the tip is short and lacks teeth laterally
(Fig. 4C’).
In species belonging to the fraterculus
group, differences were observed in the
length of aculeus tip and shape of its teeth.
The aculeus of A. /udens varies in length
from 3.2—5.0 mm and has scant broad,
rounded teeth which occupy less than the
apical half. In A. fraterculus it measures
from 1.6—1.8 mm and has large, rounded
teeth occupying half of the apical end;
while in A. obliqua it measures from 1.4—
1.7 mm and has large, sharp teeth which
occupy three-fourths of the apical end
(Figs. 4 D’—E’-F’).
Male Reproductive System
The general terminology used to describe
the male reproductive system is based on
the works of Matzuda (1976) and McAlpine
(1981), whereas for particular internal
structures we followed Hanna (1938). We
have also included a comparative synopsis
731
of the terms employed for Anastrepha by
some other authors (Table 3).
The male reproductive system is made up
of two testes, two vas deferens, an ejacu-
latory duct, two seminal vesicles, several
pairs of accessory glands, a sperm pump,
an aedeagal gland and the aedeagus itself
(Fig. 5).
The testes are oval-shaped and intensely
bright yellow. In the apical part of each tes-
tis there is a germarium followed by regions
of primary spermatocytes, secondary sper-
matocytes, spermatids, and sperm bundles,
beyond which lies the basal region which,
in mature males, contains a large quantity
of free spermatozoids (Fig. 6A). Each testis
opens into a vas deferens characterized by
a very broad anterior part which thins out
further on, finally opening into the anterior
part of the ejaculatory duct.
In the ejaculatory duct, which is very
long, three regions can be distinguished: the
widest, the anterior region, into which two
vas deferens, two seminal vesicles, and var-
ious pairs of accessory glands open; the in-
termediate region, which has a smaller di-
ameter than the first and which, extremely
thin at the end, runs through the base of the
sperm pump; and the posterior region,
which begins beyond the sperm pump and
has a smaller diameter than the former two
regions. Most of the last region is covered
by a thick pleated wall, which makes its
diameter look bigger; however, the external
wall disappears distally immediately before
reaching the basiphallus. At precisely this
point, it connects to the aedeagal gland.
The accessory glands are paired, long
and translucid; their wall is slightly thicker
than the seminal vesicles.
There are two elongated seminal vesicles
with thin walls; free spermatozoids, appar-
ently immersed in a secretion, are visible in
their interior. The vesicles open into the an-
terior part of the ejaculatory duct parallel to
the vas deferens (Fig. 6 B—C).
The sperm pump is pear-shaped with a
narrow base; it is composed of two highly
sclerotized structures of a cuticular nature.
732 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Microphotographs of the cuticular intima of the bursa copulatrix showing the ventral receptacle and
area of the vaginal duct near the signum (clorazol black E). A, A’, Anastrepha serpentina. B, B’, A. striata. C,
C’, A. cordata. D, D’, A. ludens. E, E’, A. obliqua. F, F’, A. fraterculus. Abbreviations: be = bursa copulatrix;
co = common oviduct; si = signum; vr = ventral receptacle.
VOLUME 99, NUMBER 4 WBS
Table 3. Comparison of some terminology used for the male reproductive system in the Tephritidae.
Present Study Hanna (1938) Drew (1969) Dodson (1978) Williamson Bressan (1995) Martinez et al
(1989) (1995)
Testis testis testis testis testis testis tesus
Vas deferens vas deferens vas deferens vas deferens vas deferens vas deferens vas deferens
Ejaculatory vas efferents ejaculatory common duct ejaculatory vas efferents ejaculatory
duct duct duct duct
Accessory accessory accessory accessory accessory accessory accessory
glands glands glands glands glands glands glands
Seminal vesi- end of vas de- not indicated not indicated base of testis base of testis not indicated
cles ferens
Sperm pump —_ pump organ pump organ
sperm pump
ejaculatory ejaculatory ejaculatory
apodeme apodeme apodeme
Aedeagal gland oval gland not indicated not indicated 2nd accessory not indicated
gland gland
Aedeagus aedeagus aedeagus aedeagus aedeagus aedeagus not indicated
Distiphallus not indicated not indicated
terminal end
not indicated not indicated not indicated
The ejaculatory apodeme is long and spat-
ula-shaped with a thick layer of muscles in-
serted onto it longitudinally. The other
structure is capsule-like and semi-spherical;
it surrounds the ejaculatory duct and the
ejaculatory apodeme inserts into it (Fig. 6
D). Due to the complexity of this organ, it
is very difficult to determine the exact po-
sition of the ejaculatory duct.
The aedeagal gland is an unpaired, elon-
gated, and usually voluminous organ. It is
formed by glandular units which surround
the cuticular glandular reservoir. The basal
region of the reservoir thins out slightly and
opens into a space between the wall of the
ejaculatory duct and the aedeagus, right
next to the basiphallus (Fig. 6D). This gland
has a very similar shape to that of all other
studied tephritid species.
The length of the aedeagus is variable
among the species of Anastrepha and is
probably correlated with the length of the
female terminalia in this genus (Norrbom,
personal communication). Although a sta-
tistical analysis was not performed, for
some species aedeagus length was mea-
sured and compared with average aculeus
length. We found that the species with a
long aedeagus has a long aculeus as well.
For instance, in A. fraterculus and A. obli-
qua, the aculeus is 1.4—1.8 mm long, while
the aedeagus is 2.38—2.72 mm long. In con-
trast, species with long aculeus such as A.
cordata (4.7—5.5 mm) and A. ludens (3.2—
5 mm) have an aedeagus length of approx-
imately 6.18 and 5.71 mm, respectively.
The male terminalia involves structures
such as the epandrium, proctiger and the
inner and outer surstyli. The epandrium is
a rigid and very sclerotized structure of
semi-spherical shape. In its medial posterior
region lies the membranous proctiger,
which can be retracted or expanded. The
surstyli rise out of the inferior base of the
epandrium; both the inner and outer surstyli
are partially fused together.
The most important differences among
males were observed in structures such as
the accessory glands, ejaculatory apodeme,
distiphallus and male terminalia.
In the species examined, the accessory
glands varied in number, shape and size.
Anastrepha serpentina has six pairs, A.
striata four pairs and A.cordata only two
pairs. In the species of the fraterculus group
(cf. A. ludens, A. obliqua and A. fratercu-
lus), seven pairs were observed. Forked
long glands are found in A. serpentina,
A.cordata, A. ludens, A. obliqua and A. fra-
terculus; only A. striata. has one pair of
simple long glands. Forked medium glands
were observed only in A. serpentina, A.
striata, A. obliqua and A. fraterculus. All
species, however, have simple medium
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
734
‘ajnsdeo [eoayjeuniods
— os ‘jonp yeosoujeuieds = pds ‘umrpoeyjida jesoyjeutiods = as :suoneiasiqqy ‘snjnosajosf ‘y { “pnbygo “y “y ‘suapn] ‘y ‘qd
‘pIDpAOd “YW “Dd “vIDIAIS “YW “gq “DUIJUadsas DYydasispUY “VY “(U22I13 JYSI]-uId3[NI4) sesoyeULIEds oy) Jo sydessojoydoss] “¢ “B14
VOLUME 99, NUMBER 4
Fig. 4. Outlines of the spermathecal capsules and aculeus tips. A,A’, Anastrepha serpentina. B, B’, A. striata.
C, C’, A. cordata. D, D’, A. ludens. E, E’, A. obliqua. F, F’, A. fraterculus.
glands, although in varying number: there
is one pair in A. cordata, A. obliqua, and
A. fraterculus, two pairs in A. striata and
A.ludens and four pairs in A. serpentina.
Only in A. ludens, A. obliqua and A. fra-
terculus are there four pairs of small glands.
The same number, shape and size of glands
are found only in A. obliqua and A. frater-
culus. (Fig. 7, Table 4).
In the mature males studied, the ejacu-
latory apodeme tends to be narrow in A.
ludens, A. fraterculus, and A. striata,
whereas in the others, especially A. cordata,
is visibly wider at its apical half (Fig. 8).
However, the differences should be consid-
ered with some reserve, due to the fact that
intraspecific size can vary depending on the
degree of maturity of each individual (Drew
1969). To study this properly, careful com-
parison must be made of each species over
time, and the results presented should in-
clude a range of intraspecific variation.
In general, the distiphallus (Fig. 9) has a
bulky basal portion known as the basal
lobe, with tiny microspines on its surface.
The rest is a membranous, semitransparent
unit with an interior sclerotized duct. All
Anastrepha and Toxotrypana species pre-
sent an internal apical sclerite “‘T’’ shaped
(sensu Norrbom 1985).
Within the male terminalia, some inter-
specific differences stand out, particularly
the shape of the outer surstyli when viewed
laterally. In most of the studied species,
they were elongated, except in A. cordata
in which the surstyli are very short and
widened at the base (Fig. 10).
DISCUSSION
The general anatomy of the female re-
productive system is similar in all of the
studied Anastrepha species, as well as other
Tephritidae such as Rhagoletis pomonella
(Dean 1935), C. capitata (Hanna 1938), B.
736
eae
Fig. 5. General morphology of the reproductive sys-
tem in the male of Anastrepha serpentina. Abbrevia-
tions: aeg = aedeagal gland; ag = accessory gland; bp
= basiphallus; eae = entrance of the ejaculatory duct
to aedeagus; ed = ejaculatory duct; spp = sperm
pump; sv = seminal vesicles; te = testis; vd = vas
deferens.
tryoni (Drew 1969) and B. oleae Gmelin
(Solinas and Nuzzaci 1984).
The number of ovarioles present in the
Six species of Anastrepha varies between
20 and 33 per ovary. This number is also
very similar to that of other species of fruit
flies such as C. capitata, which has an av-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
erage of 28 ovarioles (Hanna 1938), R. po-
monella with an average of 24 (Dean 1935),
and Bactrocera tryoni with about 22 (An-
derson and Lyford 1965). However, species
such as R. conversa and R. nova have an
average of 9 and 12 ovarioles, respectively
(Flores et al. 1987).
In all Anastrepha females, the ovarian
follicle contains the oocyte and 15 nutritive
cells, similar results were reported previ-
ously for A. obliqua (Bressan and Costa Te-
les 1991) and other species such as B.
tryoni (Anderson and Lyford 1965), R. con-
versa and R. nova (Flores et al. 1987).
The anatomy of the vagina in the Te-
phritidae, except in the case of R. pomo-
nella (Dean 1935), has never been clearly
described. In C. capitata the vagina and
vaginal duct were depicted without any
mention of the bursa copulatrix and the
ventral receptacle (Hanna 1938, Guillén
1983). The ventral receptacle was illustrat-
ed as an unpaired, sclerotized organ in X.
unipuncta by Souza Lopes (1939) as well
as in other groups of Tephritidae (Munro
1984). In B. tryoni this structure was termed
the morula gland by Drew (1969).
In the few species of Anastrepha studied
previously, neither the bursa copulatrix nor
the ventral receptacle has been adequately
described. With respect to A. suspensa and
A. ludens, the posterior region of the vagina
was mislabeled the oviduct by Dodson
(1978), while Servin-Villegas and Jiménez-
Jiménez (1995) believed that the vagina is
formed by a widening in the oviduct. In A.
Table 4. Comparison of some characteristics of the accessory glands of males of six Anastrepha species.
Long
Species Forked
. Serpentina
. striata
. cordata
. ludens
. obliqua
. fraterculus
baths HS bs Ths
meee |e
|
Accessory glands (pairs)
Medium
Forked Simple Small Total
1 4 — 6
1 2 ms 4
= l ~_ p:
= 2 4 f
1 1 4 7
1 1 4 7
VOLUME 99, NUMBER 4 U3
‘“d - q200 um
wa: “we J
~~ *
Fig. 6. Microphotographs of some structures of the reproductive system in the male of Anastrepha serpentina
(Feulgen-light green). A, Testis. B, Accessory glands. C, Seminal vesicles and vas deferens. D, Sperm pump
and aedeagal gland. Abbreviations: aeg = aedeagal gland; ag = accessory glands; bp = basiphallus; sb = sperm
bundles; spp = sperm pump; sv = seminal vesicles; sz = spermatozoids; vd = vas deferens.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 7. Outlines of the accessory glands and seminal vesicles. A, Anastrepha serpentina. B, A. striata. C, A.
cordata. D, A. ludens. E, A. obliqua. F, A. fraterculus. Abbreviations: ag = accessory glands; ed = ejaculatory
duct; sv = seminal vesicles; vd = vas deferens.
serpentina, only the anterior vagina and
vaginal duct have been recognized (Marti-
nez et al. 1995).
Other authors such as Dean (1935) and
Williamson (1989) characterize the vagina
as consisting of two regions: the anterior,
which is made up of the bursa copulatrix
as well as the ventral receptacle, also called
the fertilization chamber by Solinas and
Nuzzaci (1984). The second is the posterior
region, which consists of the vaginal duct.
In insects the bursa copulatrix acts as a
receptacle for the male intromittent organ
(Snodgrass 1935). In Anastrepha it receives
the distiphallus during copula, but sperma-
tozoids are deposited in the ventral recep-
tacle before being transferred to the sper-
mathecae. The few which remain in the
ventral receptacle will be the first sperma-
tozoids to fertilize the eggs (Dean 1935, So-
linas and Nuzzaci 1984).
The presence of three spermathecae in all
species of Anastrepha is characteristic of
most Trypetinae (sensu Hancock 1986a),
except in some species of Rhagoletis such
as R, nova and R. conversa (Flores et al.
1987), R.striatella Wulp (Bush 1966) and
other Carpomyina (Norrbom 1994). In con-
trast, Dacinae species such as C. capitata,
B. tryoni and B. oleae have only two sper-
mathecae (Hanna 1938, Drew 1969, Solinas
and Nuzzaci 1984), as well as all Tephriti-
nae (Hancock 1986b).
Female accessory glands in tephritid fe-
males have been misrepresented as colle-
terial glands by some authors (Hanna 1938,
Drew 1968, Flores et al. 1987). This term
is employed specifically for the accessory
glands of the Orthoptera, in which the col-
leterial glands are responsible for secreting
substances which form the ootheca (Davey
1985a). In Diptera, the function of the ac-
cessory glands is poorly understood, except
in Glossina (Glossinidae), in which they are
called milk glands because their secretions
serve as nourishment for the intrauterine
larvae (Matzuda 1976, Davey 1985a).
For tephritids, the function of the acces-
sory glands is not well understood, al-
though Christenson and Foote (1960) spec-
ulated that their secretions act as a vaginal
lubricant before oviposition. Solinas and
Nuzzaci (1984) mentioned that these secre-
tions can help carry sperm toward the fer-
tilization chamber. It has also been sug-
gested that the accessory glands produce
VOLUME 99, NUMBER 4
: i mi A
Fig. 8. Microphotographs of ejaculatory apodemes of the sperm pump (chlorazol black E). A, Anastrepha
serpentina. B, A. striata. C, A. cordata. D, A. ludens. E, A. obliqua. F, A. fraterculus.
marking pheromones deposited by the fe-
male after oviposition so that other females
recognize infested fruit (Prokopy and Roit-
berg 1984).
The general anatomy of the male repro-
ductive system for Anastrepha is similar in
all studied species as well as other Tephrit-
idae (Hanna 1938, Souza Lopes 1939,
Drew 1969).
As to the location of the seminal vesicles
in Tephritidae, various authors (Anwar
1971, Williamson 1989, Bressan 1995, Ser-
vin-Villegas and Jiménez-Jiménez 1995)
stated that they are located to the basal part
of the testis perhaps due to the abundant
presence of free spermatozoids. In C. cap-
itata they confused the seminal vesicles
with the anterior part of the ejaculatory duct
(Hanna 1938). As to X. unipunctata, Souza
Lopes (1939) stated that the seminal vesi-
cles are two structures which open into the
anterior region of the ejaculatory duct near
the vas deferens.
In mature males of Anastrepha, free sper-
740
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 9. Outlines of distiphallus. A, Anastrepha serpentina. B, A. striata. C, A. cordata. D, A. ludens. E, A.
obliqua. FE A. fraterculus.
matozoids were observed at the base of the
testes, all along the vas deferens and in the
seminal vesicles. In this genus the seminal
vesicles open into the ejaculatory duct par-
allel to the vas deferens and between the
accessory glands, as is the case in other
groups of Diptera (see Matzuda 1976). In
Anastrepha the seminal vesicles have been
confused with the accessory glands due to
their proximity to them and because no
spermatozoids have been observed in the
seminal vesicles.Their function for the stor-
age of the mature spermatozoids until the
moment of copula is well known in differ-
ent species of insects (Snodgrass 1935,
Davey 1985b).
The accessory glands of Anastrepha
males has not been described in detail pre-
viously. It is quite probable that they are
responsible for secreting substances which
form the seminal liquid which accompany
the spermatozoids; this has been docu-
mented for other groups of insects (Davey
1985b). The species examined from the fra-
terculus group all had seven pairs of acces-
sory glands, the highest number observed.
The other species had from two to six pairs,
and the number differing from species to
species (Table 3). In A. suspensa, which
also belongs to the fraterculus group, the
existence of three to four pairs was men-
tioned by Dodson (1978), It is probable that
this author did not observe the four pairs of
small glands because they are difficult to
discern without dye; he may have mistaken
one pair for seminal vesicles. For A. /udens,
Servin-Villegas and Jiménez-Jiménez
(1995) reported the presence of four pairs,
which in all likelihood correspond to the
pair of long glands, two pairs of medium
glands and one pair of seminal vesicles.
The four small glands were not observed
(Table 4).
Xanthaciura unipuncta has only two
pairs of accessory glands (Souza Lopes
1939). For C. capitata, Hanna (1938) and
Valdés Carrasco and Prado Beltran (1990)
described the presence of four pairs, one
very long and three short, although we as-
sume that one pair is actually the seminal
vesicles. For B. tryoni, Drew (1969) de-
scribed 4 pairs, but one of them also exhib-
its marked morphological differences which
VOLUME 99, NUMBER 4
741
0.3 mm
Fig. 10. Simplified outlines of the male genitalia in lateral view. A, Anastrepha serpentina. B, A. striata. C,
A. cordata. D, A. ludens. E, A. obliqua. F, A. fraterculus. Abbreviations: ep = epandrium; pr = proctiger; ss =
outer surstyl1.
make us believe that they are seminal ves-
icles.
The sperm pump is made up of a semi-
spherical base to which a sclerotized inter-
nal structure called the ejaculatory apodeme
is attached; many muscles are inserted onto
the apodeme. The sperm pump pumps sem-
inal liquid and distends the aedeagus during
copula (Matzuda 1976).
In this respect, most studies of Tephriti-
dae call the sperm pump the ejaculatory
apodeme, while other authors such as Han-
na (1938) and Drew (1969) call it the erect-
ing and pumping organ.
In all Anastrepha species examined, the
aedeagal gland corresponds to the oval-
shaped gland described for C. capitata by
Hanna (1938), and to the second accessory
gland indicated by Bressan (1995) for
Anastrepha spp. Given its glandular ana-
tomical characteristics and because it opens
into a region between the ejaculatory duct
and the aedeagus at the base of the basi-
phallus, its secretions must empty into this
space. The secretions may function as a lu-
bricant, as suggested by Hanna (1938), and
may also serve to distend the aedeagus dur-
ing copula.
ACKNOWLEDGMENTS
This study was carried out with the sup-
port of the Instituto de Ecologia (Ref: 902—
38). We thank Gonzalo Pérez Higareda, Di-
rector of the “‘Estacién de Biologia Tropical
Los Tuxtlas,” UNAM, for providing us
with the facilities to collect study material.
742
We are also grateful to our colleagues Mar-
tin Aluja S. and Isabel Jacome A. for pro-
viding food resources for the adults reared
in the laboratory and for part of the material
examined. We thank Magdalena Cruz R. for
her valuable support in creating the illustra-
tions and Ingrid Crews for translating the
manuscript into English. We are also grate-
ful to Dr. A.L. Norrbom, Systematic Ento-
mology Laboratory, USDA, and an anony-
mous reviewer for their constructive criti-
cism and suggestions.
LITERATURE CITED
Anderson, D. T. and G. C. Lyford. 1965. Oogenesis
in Dacus tryoni (Frogg) (Diptera: Trypetidae).
Australian Journal of Zoology 13:423—435.
Anwar, M., D. L. Chambers, K. Ohinata and R. M.
Kobayashi. 1971. Radiation sterilization of the
Mediterranean fruit fly (Diptera: Tephritidae);
Comparison of spermatogenesis in flies treated as
pupae or adults. Annals of the Entomological So-
ciety of America 64:627—633.
Bressan, S. 1995. Desenvolvimento e potencial repro-
dutivo do macho de Anastrepha spp. (Diptera: Te-
phritidae) em condicoes naturais. Revista Brasi-
leira de Entomologia 39(4):849-854.
Bressan, S. and M. Costa Teles. 1991. Ovogénese em
Anastrepha obliqua (Macquart) (Diptera, Tephrit-
idae). Revista Brasileira de Entomologia 35(4):
691-696.
Bush, G. L. 1966. The taxonomy, cytology and evo-
lution of the genus Rhagoletis in North America
(Diptera, Tephritidae). Bulletin of the Museum of
Comparative Zoology 134(11):431—560.
Carayon, J. 1969. Emploi du noir chlorazol en ana-
tomie microscopique des insectes. Annales de la
Société Entomologique de France (N.S.) 5(1):
179-193.
Christenson, L. D. and R. H Foote. 1960. Biology of
fruit flies. Annual Review of Entomology 5:171—
192%
Dampf, A. 1933. Estudio sobre el oviscapto de las
moscas de la fruta (Anastrepha spp.) de México.
Irrigaci6n en México (Comisié6n Nacional de Ir-
rigacion de la Secretaria de Agricultura y Fo-
mento) 6(3):253—265.
Davey, K. G. 1985a. The female reproductive tract,
pp. 15-36. Jn Kerkut G. A. and L.I. Gilbert, eds.
Comprehensive Insect physiology, biochemistry
and pharmacology. Pergamon Press. Vol.I.
Davey, K. G. 1985b. The male reproductive tract, pp.
1-14. Jn Kerkut G. A. and L.I. Gilbert, eds. Com-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
prehensive Insect physiology, biochemistry and
pharmacology. Pergamon Press. Vol.I.
Dean, R. W. 1935. Anatomy and postpupal develop-
ment of the female reproductive system in the ap-
ple maggot fly, Rhagoletis pomonella Walsh. New
York State Agricultural Experimental Station
Technical Bulletin 229:1—31.
Dodson, G. 1978. Morphology of the reproductive
system in Anastrepha suspensa (Loew) and notes
on related species. The Florida Entomologist
61(4):231-239.
Drew, R. A. I. 1969. Morphology of the reproductive
system of Strumeta tryoni (Froggatt) (Diptera:
Trypetidae) with a method of distinguishing sex-
ually mature adult males. Journal of the Australian
Entomological Society 8:21—32.
Flores, R. V., D. Frias and A. H. Martinez. 1987. His-
tologia y morfologia del sistema reproductor en
hembras de Rhagoletis conversa (Brethes) y de
Rhagoletis nova (Schiner) (Diptera, Tephritidae).
Revista Brasileira de Zoologia 4(1):33—46.
Gabe, M. 1968. Techniques histologuiques. Ed. Mas-
soneu Cie iS app:
Guillén, A. J. C. 1983. Manual para la diferenciacién
de moscas del mediterraneo Ceratitis capitata
(Wied.) silvestres (fértiles) de moscas irradiadas
(estériles). Programa Mosca del Mediterraneo,
SARH. 102 pp.
Hancock, D. 1986a. Classification of the Trypetinae
(Diptera: Tephritidae) with a discussion of the Af-
rotropical fauna. Journal of Entomological Society
of South Africa 49:275—305.
Hancock, D. 1986b. New genera and species of Af-
rican Tephritinae (Diptera=phritiade), with com-
ments on some currently unplaced of misplaced
taxa and on classification. Transactions of the
Zimbabwe Scientific Association 63:16—34.
Hanna, A. D. 1938. Studies on the Mediterranean fruit
fly: Ceratitis capitata (Wied.) I. The structure and
operation of the reproductive organs. Bulletin de
la Societé Foread Ier De Entomologie 22:39—S2.
Hernandez-Ortiz, V. and M. Aluja. 1993. Listado de
especies del género neotropical Anastrepha (Dip-
tera: Tephritidae) con notas sobre su distribucién
y plantas hospederas. Folia Entomologica Mexi-
cana 88:89-105.
Martinez, M. I., V. Hernandez-Ortiz and L. R. Luna.
1995. Desarrollo sexual en Anastrepha serpentina
(Wiedemann) (Diptera: Tephritidae). Acta Zool6-
gica Mexicana (n.s.) 65:75-88.
Matzuda, R. 1976. Morphology and evolution of the
insect abdomen. Pergamon, Press. pp. 335-363.
McAlpine J. FE 1981. Morphology and terminology—
adults, pp. 9-63. In McAlpine J. F, B. V. Peter-
son, G. E. Shewell, H. J. Teskey, J. R. Vockeroth
and D. M. Wood, eds. Manual of Nearctic Diptera.
Research Branch Agriculture Canada. Monograph
Pile
VOLUME 99, NUMBER 4
Munro, H. K. 1984. A taxonomic treatise on the Dac-
idae (Tephritoidea, Diptera) of Africa. Entomolo-
gy Memoirs. Department of Agriculture and Water
Supply. Republic of South Africa. 61, ix + 1-313.
Norrbom, A. L. 1985. Phylogenetic analysis and tax-
onomy of the cryptostrepha, daciformis, robusta
and schausi species groups of Anastrepha Schiner
(Diptera: Tephritidae). Pennsylvania State Univer-
sity (Ph.D. Thesis). 355 pp.
Norrbom, A. L. and K. C. Kim. 1988. Revision of the
schausit Group of Anastrepha Schiner (Diptera,
Tephritidae), with a Discussion of the Terminol-
ogy of the female terminalia in the Tephritoidea.
Annals of the Entomological Society of America
81(2):164-173.
Norrbom, A. L. 1994. New species and phylogenetic
analysis of Cryptodacus, Haywardina, and Rhag-
oletotrypeta (Diptera: Tephritidae). Insecta Mundi
8(1—2):37-65.
Prokopy, R. J. and B. D. Roitberg. 1984. Foraging
behavior of true fruit flies. American Scientist 72:
41-49.
Servin-Villegas, R. and M. L. Jiménez-Jiménez. 1995.
743
El aparato reproductor y ciclo gonadico de la mos-
ca de la fruta Anastrepha ludens. Southwestern
Entomologist 20(2):223-227.
Snodgrass, R. E. 1935. Principles of insect morphol-
ogy. McGraw-Hill Book Company. 667 pp.
Solinas, M. and G. Nuzzaci. 1984. Functional anato-
my of Dacus oleae Gmel. female genitalia in re-
lation to insemination and fertlization processes.
Entomologica, XIX Bari 16(12):135—165.
Souza Lopes, H. de. 1939. Sobre a Xanthaciura uni-
puncta Malloch, 1933 (Dipt.: Trypetidae), pp.
424-431. In Livro de Homenagem aos Prof. Al-
varo e Miguel Ozorio de Almeida, Rio de Janeiro,
Brasil.
Valdés-Carrasco, J. and E. Prado-Beltran. 1990. Es-
queleto y musculatura de la mosca del mediter-
raneo, Ceratitis capitata (Wiedemann) (Diptera:
Tephritidae). Folia Entomologica Mexicana 80:
59-225.
Williamson, D. L. 1989. Oogenesis and spermatogen-
esis, pp. 141-151. Jn Robinson A. S. and Hooper,
G., eds., Fruit flies. Their biology, natural enemies
and control Vol. 3A.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 744-747
AIVALYKUS DOMINICANUS (HYMENOPTERA: BRACONIDAE), A NEW
SPECIES FROM DOMINICAN AMBER
ROBERT L. ZUPARKO AND GEORGE O. POINAR, JR.
(RLZ) Essig Museum of Entomology, 201 Wellman Hall, University of California,
Berkeley, CA 94720, U.S.A.; (GOP) Department of Entomology, Oregon State University,
Corvallis, OR 97331, U.S.A.
Abstract.—Alvalykus dominicanus, n. sp., is described from two male specimens pre-
served in Dominican amber, and represents the first described species of the genus from
the Neotropics. The specimens are 15—45 million years old. The new species is remarkable
by the elongated abdominal terga (longer than the rest of the body), and is distinguished
from the four extant Aivalykus species by a more elongated first abdominal tergite and
differences in fore wing venation.
Key Words:
The genus Aivalykus Nixon 1938 (Hy-
menoptera: Braconidae) is placed in the
Doryctinae, tribe Hecabolini, and can be
distinguished from other genera of the tribe
by the following combination of characters:
males with elongated abdominal terga, fem-
ora not swollen, fore wing with nervulus
present, stigmal length greater than 2 times
width, radius reaching wing margin, dis-
coideus and subdiscoideus interstitial. The
genus contains four extant species from the
Indo-Pacitic, Malagasy and Nearctic
regions. Males are known from only two
species, A. eclectes Nixon (the genotype)
and A. nearcticus Marsh, that are remark-
able for their elongated abdomens (Nixon
1938, Marsh 1965). We have examined two
male Aivalykus specimens imbedded in Do-
minican amber. These represent a new spe-
cies, which is described below.
These specimens originated from mines
located in the Cordillera Septentrional, be-
tween Santiago and Puerto Plata, in the
northern portion of the Dominican Repub-
lic. These mines are in the El Mamey For-
mation (Upper Eocene), which is a shale-
sandstone interspersed with a conglomerate
Braconidae, Aivalykus dominicanus n. sp., fossils, Dominican amber
of well-rounded pebbles (Eberle et al.
1980). The exact age of the amber is un-
known, but estimates based on various mi-
crofossils and chemical analyses provide a
range from 15-20 million years (Iturralde-
Vinent and MacPhee 1996) to 30—45 mil-
lion years (Cepek in Schlee 1990).
Aivalykus is the fourth genus (along with
Hecabolus Haliday, Polystenus Forster and
Promonolexis Brues, all known from Eu-
ropean Oligocene specimens) in the Heca-
bolini represented by fossil remains; addi-
tional doryctinine genera with recorded fos-
sils include Ecphylus Forster (from Mexi-
can amber, 22—26 mya), Doryctes Haliday
(from Baltic amber, 40 mya) and Rhaco-
notus Ruthe (from Florissant beds, 34 mya)
(Carpenter 1992).
Terminology of wing venation follows
Marsh et al. (1987). Specimens were stud-
ied immersed in corn oil.
Aivalykus dominicanus Zuparko and
Poinar, NEW SPECIES
(Figs. 1-3)
Diagnosis.—Length of first abdominal
tergite 2.5 times its apical width, and re-
VOLUME 99, NUMBER 4
Figs. 1-3.
3, Fore wings.
current vein of fore wing received into Ist
cubital cell by a distance about %4 its own
length. The four other described Aivalykus
species have length of first tergite less than
2.5 times apical width; recurrent vein of A.
eclectes is interstitial or received into 2nd
cubital cell, that of A. niger Granger into
the 2nd cubital cell, and that of A. nearc-
ticus received into Ist cubital cell by a dis-
tance equal to about % its length. Addition-
ally, fore wing stigma of A. dominicanus is
less than 4 times its width (greater than 4
times its width in A. niger), in profile dor-
sum of mesonotum very flat (rounded in A.
nearcticus), and the sternaulus extends over
the greater part of the mesopleuron (absent
in A. sperches Nixon).
Description.—Male (habitus; Fig. 1):
745
Aivalykus dominicanus (male). 1, Habitus (Scale bar = 1.0 mm). 2, Dorsal aspect of antennae.
overall length (inclusive of abdominal ter-
ga), 3.5-4.0 mm; body length (exclusive of
abdominal terga posterior to last abdominal
sternum), 1.5 mm. Apparently bronze, with
scape, central portion of frons, propleuron,
ventral portion of mesopleuron, propo-
deum, central portion of anterior sterna and
legs (except tarsi), darkened. Head about
twice as wide as long, narrower behind eyes
than across vertex, vertex finely reticulate.
Relative length: width of antennal seg-
ments: scape—4 X 2.5, pedicel—3 X 2,
first funicular segment (Fl)—8 Xx 1, F2—
9 X 1, remaining funicle segments—6—7
1 (antennae broken after F1l1); from dorsal
apsect, Fl medially curved markedly in-
ward, F2 curved slightly less so (Fig. 2).
Mesosoma approximately equal in length
746
to metasoma (exclusive of projecting terga
which are about 3 X its length). Apical
margin of pronotum with short upturned
transparent lamella. In profile, mesonotum
anteriorly declining almost perpendicularly
to pronotum, remainder of mesonotum, scu-
tellum and anterior part of propodeum very
flat, laying along one plane, posterior part
of propodeum sloping down at an angle of
about 30°. Small lateral dorsal tubercle on
mesonotum at junction of longitudinal and
perpendicular planes; notauli distinct, con-
verging posteriorly; mesonotum finely re-
ticulate. Mesopleuron smooth, sternaulus
extending about 70% length of sclerite. No
indication of metanotal spine, propodeum
appearing smooth. Fore coxa globular, mid
coxa about 1.5 times longer than wide, hind
coxa about 2 times longer than wide; fem-
ora not swollen, Ist segment of tarsi about
twice length of 2nd segment on all legs.
Fore wing (Fig. 3): recurrent vein entering
first cubital cell by a distance about % its
own length; length of stigma less than 4
times its width; parastigma appearing as a
widening of the basal vein, extending from
the stigma about % the distance to the cu-
bitus; 2nd abcissa of radius slightly up-
curved, forming an obtuse angle of about
130° with first abcissa; nervulus faint, en-
tering Ist discoidal cell; cubitus strongly
pigmented for about 40% its length past the
Ist intercubitus, very lightly pigmented
thereafter; Ist brachial cell open; subdis-
coideus strongly pigmented for about 30%
its length past Ist recurrent vein, lightly
pigmented thereafter.
Length of first abdominal tergum about
2.5 times its width, widening slightly pos-
teriorly; smooth over most of its surface,
slightly excavated posteriorly with short
longitudinal striations posteriorlaterally.
Second abdominal tergum about 2 times
long as wide, narrowing slightly posterior-
ly. All succeeding terga several times lon-
ger than wide, more or less parallel sided,
terminal tergum triangular, narrowing api-
cally. Terminal sternum extending slightly
past second tergum.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Female—Unknown.
Material examined—Holotype, a well-
preserved male in a dark yellow piece of
amber, 15 X 7 X 7 mm, number H-10-93.
From Cordillera Septentrional, between
Santiago and Puerto Plata, in the Domini-
can Republic. Deposited in the Poinar am-
ber collection maintained at Oregon State
University, Corvallis, Oregon.
Paratype, | d, in very poor condition, in a
very light yellow piece of amber, 20 X 11
x 7 mm, number H-10-93A. Collection and
deposition data same as holotype.
Etymology—tThe specific epithet domini-
canus, a neolatin noun in apposition, refers
to the Dominican Republic, the country of
origin of the specimens.
DISCUSSION
Except for the terminal antennal seg-
ments, the holotype is virtually complete
and viewable from several angles. The
paratype is in extremely poor condition and
is recognizable solely by the flattened ex-
tended abdominal terga.
Of the four extant Aivalykus species, two
(A. niger Granger and A. sperches Nixon)
have unknown hosts, and are described
only from female specimens. The other two
species (A. eclectes and A. nearcticus) are
associated with bark beetles (Coleoptera:
Scolytidae) and characterized by sexually
dimorphic abdominal shapes (Beeson 1941,
Marsh 1965). Like A. dominicanus, the
males have greatly elongated abdominal
terga, though not to the same degree. In all
four extant species the females do not have
excessively elongated abdomens.
Elongated ovipositors and/or abdomens
are common characteristics of female par-
asitoids which oviposit in or on hosts (such
as bark beetles) hidden deep in a substrate,
while the conspecific males often have
shorter abdomens. Thus the reversal seen in
Aivalykus (females with short abdomens
and males with long abdomens) is highly
unusual. This condition may reflect a male
mating-behavior strategy to be the first to
mate with emerging females. Similar be-
VOLUME 99, NUMBER 4
havior has been found in the Rhyssini (Hy-
menoptera: Ichneumonidae—parasitoids of
deep-boring siricoid wood wasps). Males of
Megarhyssa and Rhyssella (which have
bendable abdomens) may increase their re-
productive success by being able to remain
on the surface of the bark and mate with
females before the latter leave the emer-
gence burrow (Godfray 1994). In the case
of Aivalykus, we theorize the elongated
male abdomen reflects the distance the male
needs to cover in order to contact the fe-
male still in the host gallery. Although one
would expect to find a similarly-lengthened
abdomen in the female in order to oviposit
to the same depth, this is in fact not a nec-
essary condition in bark beetle parasitoids.
Roptrocerus xylophagorum (Ratzeburg)
(Hymenoptera: Pteromalidae) and Entedon
leucogramma (Ratzeburg) enter bark beetle
galleries to oviposit (Reid 1957, Beaver
1966), while Tomicobia tibialis Ashmead
(Hymenoptera: Pteromalidae) oviposits on
adult bark beetles before they enter the gal-
leries (Reid 1957).
ACKNOWLEDGMENTS
We are grateful to P Marsh (North New-
ton, Kansas) for confirming the generic
identity of A. dominicanus. We also thank
S. Heydon (University of California, Davis)
for the loan of A. nearcticus paratypes for
study, and Mike Sharkey (University of
Kentucky, Lexington), and one anonymous
reviewer for their useful comments.
747
LITERATURE CITED
Beaver, R. A. 1966. The biology and immature stages
of Entedon leucogramma (Ratzeburg) (Hymenop-
tera: Eulophidae), a parasite of bark beetles. Pro-
ceedings of the Royal Entomological Society of
London (A) 41: 37-41.
Beeson, C. EF C. 1941. The ecology and control of the
forest insects of India and the neighboring coun-
tries. The Author, Dehra Dun, India. 1007 pp.
Carpenter, F M. 1992. Part R: Arthropoda, Vol. 4:
Superclass Hexapoda. /n Treatise on Invertebrate
Paleontology, Geological Society of America and
University of Kansas, Lawrence, Kansas.
Eberle, W., W. Hirdes, R. Muff, and M. Pelaez. 1980.
The geology of the Cordillera Septentrional. Pro-
ceedings of the 9th Caribbean Geological Confer-
ence, August 1980, Santo Domingo, pp. 619-632.
Godfray, H. C. J. 1994. Parasitoids: Behavioral and
Evolutionary Ecology. Princeton University Press,
Princeton, New Jersey. 473 pp.
Iturralde-Vinent, M. A. and R. D. E. MacPhee. 1996.
Age and paleogeographical origin of Dominican
amber. Science 273: 1850-1852.
Marsh, P. M. 1965. The Nearctic Doryctinae. I. A
review of the subfamily with a taxonomic revision
of the tribe Hecabolini (Hymenoptera: Braconi-
dae). Annals of the Entomological Society of
America 58: 668-699.
Marsh, P. M., S. R. Shaw, and R. A. Wharton. 1987.
An identification manual for the North American
genera of the family Braconidae (Hymenoptera).
Memoirs of the Entomological Society of Wash-
ington 13, 98 pp.
Nixon, G. E. J. 1938. A new genus of Hecabolinae
and a note on the genus Telebolus Marshall
(Hym., Braconidae). Proceedings of the Royal En-
tomological Society of London (B) 7: 152-156.
Reid, R. W. 1957. The bark beetle complex associated
with lodgepole pine slash in Alberta. Part I]—
Notes on the biologies of several hymenopterous
parasites. The Canadian Entomologist 89: 5-8.
Schlee, D. 1990. Das Bernstein-Kabinett. Stuttgarter
Beitrager fiir Naturkunde, Ser. C., No. 28: 1-100.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 748-755
LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF
TRUPANEA SIGNATA FOOTE (DIPTERA: TEPHRITIDAE) ON GNAPHALIUM
LUTEO-ALBUM L. IN SOUTHERN CALIFORNIA
RICHARD D. GOEDEN AND JEFFREY A. TEERINK
Department of Entomology, University of California, Riverside, CA 92521, U.S.A.
Abstract.—Trupanea signata L. is a narrowly oligophagous, multivoltine, gregarious,
obligately gallicolous fruit fly (Diptera: Tephritidae) studied in southern California on
Gnaphalium luteo-album L. This host plant is an introduced weedy annual of Old World
origin to which this tephritid has transferred from indigenous, congeneric hosts, an un-
common occurrence among nonfrugivorous North American Tephritidae. The egg, third-
instar larva, and puparium are described and figured. The eggs are inserted into apical
and axillary buds. From one to six larvae feed gregariously within an open cavity in the
expanded pith parenchyma of galls on axillary branches and stems during all three instars.
The galls are elongate-obclavoidal, sometimes bear axillary branches and flower heads,
and are covered by a wooly investiture. Pupariation occurs inside the gall. The adults
emerge and exit the gall through a common channel by pushing aside a thin, pre-formed,
apical, ‘‘gall cap.”’ The adults are long-lived and are the overwintering stage. The life
cycle follows the aggregative pattern and at least two generations, one each in spring and
fall, are produced annually on G. luteo-album. Two species of solitary, larval-pupal, hy-
menopterous endoparasitoids are reported: Eurytoma sp. (Eurytomidae) and Pteromalus
sp. (Pteromalidae).
Key Words: Insecta, Trupanea, Gnaphalium, biology, taxonomy of immature stages,
galls, oligophagy, parasitoids
The genus 7rupanea (Diptera: Tephriti-
dae) occurs worldwide and is a numerically
large and widespread taxon of nonfrugivo-
rous fruit flies in North America and Cali-
fornia (Foote and Blanc 1963, Foote et al.
1993). As such, the adults are among the
most commonly encountered, reared or
swept tephritids; however, the life histories
of most species remain littlhe known, and
several species are difficult to distinguish
morphologically (Foote 1960, Foote et al.
1993). Detailed life histories of five species
of Trupanea from southern California have
been published (Cavender and Goeden
1982, Goeden 1987, 1988, Headrick and
Goeden 1991, Knio et al. 1996b), and the
immature stages of three of these species
also described (Cavender and Goeden 1982,
Headrick and Goeden 1991, Knio et al.
1996a). This and our next five papers will
more than double the number of species of
Trupanea for which life histories and im-
mature stages are well known, beginning
here with the indigenous, obligate gall-for-
mer, 7. signata Foote.
MATERIALS AND METHODS
Our field studies on T. signata mainly fo-
cused on laboratory dissections of galls col-
lected on Gnaphalium luteo-album L. at
different locations in central and southern
California during 1989-91: (1) N end of
VOLUME 99, NUMBER 4
Hemet Lake at 1424-m elevation, San Ber-
nardino Nat. Forest (S section), Riverside
Co., 29.ix.1989 and 11.ix.1990; (2) along
Deer Creek in the Morton Flat area at 550
m. Tulare Co., 14.vi.1990; (3) Box Springs
Grade between Moreno Valley and River-
side at 470 m, Riverside Co., 20-
28.11.1991. Excised galls and uprooted gall-
bearing shoots were transported in cold-
chests in an air-conditioned vehicle to the
laboratory and stored under refrigeration
for subsequent dissection, photography, de-
scription, and measurement. Fifteen eggs
recovered from ovipositional cagings and
two second- and eight third-instar larvae,
and four puparia dissected from galls were
preserved in 70% EtOH for scanning elec-
tron microscopy (SEM). Additional puparia
were placed in separate, glass shell vials
stoppered with absorbant cotton and held in
humidity chambers at room temperature for
adult and parasitoid emergence. Specimens
for SEM were hydrated to distilled water in
a decreasing series of acidulated EtOH.
They were osmicated for 24 h, dehydrated
through an increasing series of acidulated
EtOH and two, 1-h immersions in Hexa-
methlydisilazane (HMDS), mounted on
stubs, sputter-coated with a gold-palladium
alloy, and studied with a JEOL JSM C-35
SEM in the Department of Nematology,
University of California, Riverside.
Most adults reared from isolated puparia
were individually caged in 850-ml, clear-
plastic, screened-top cages with a cotton
wick and basal water reservoir and provi-
sioned with a strip of paper toweling im-
pregnated with yeast hydrolyzate and su-
crose. These cages were used for longevity
studies, and those with the wicks wrapped
around root-bearing shoots of G. luteo-al-
bum for oviposition studies, in the insectary
of the Department of Entomology, Univer-
sity of California, Riverside, at 25 + 1°C,
and 14/10 (L/D) photoperiod. Virgin male
and female flies obtained from emergence
vials were paired (n = 16) in clear-plastic
petri dishes provisioned with a flattened,
water-moistened pad of absorbant cotton
749
spotted with honey (Headrick and Goeden
1991, 1994) for observations of their court-
ship and copulation behavior.
Plant names used in this paper follow
Munz (1974, as updated by Hickman
1993); tephritid names and adult terminol-
ogy follow Foote et al. (1993). Terminology
and telegraphic format used to describe the
immature stages follow Knio et al. (1996a)
and Goeden and Teerink (1996a, b, c,
1997a, b) and our earlier works cited there-
in. Means + SE are used throughout this
paper. Voucher specimens of 7. signata and
its parasitoids reside in the research collec-
tions of RDG; preserved specimens of eggs,
larvae and puparia are stored in a separate
collection of immature Tephritidae main-
tained by JAT.
RESULTS AND DISCUSSION
TAXONOMY
Adult.—Trupanea signata was described
from reared and swept adults collected from
various locations in California by Foote
(1960), who also pictured the wing of a fe-
male. Foote et al. (1993) illustrated the head
of an adult in side view and the wing pat-
tern of a female, and noted that the wing
pattern of the male *. . .does not differ from
that of the female in any important re-
spect.”’
Immature stages.—Egg: Twenty-one
eggs of T. signata were white, opaque,
smooth; with an elongate-ellipsoidal body,
0.69 + 0.005 (range, 0.65—0.74) mm long,
0.21 + 0.004 (range, 0.17—0.23) mm wide,
smoothly rounded at tapered posterior end,
and with a peg-like anterior pedicel, 0.02
mm long (Fig. 1); a single row of aeropyles
circumscribes the pedicel (Fig. 1).
The egg of T. signata is similar in shape
to the eggs of other Trupanea species pre-
viously described. The egg is larger in
width and length than 7. californica Mal-
loch, approximately the same size as T. im-
perfecta (Coquillett), and shorter than T.
conjuncta (Adams), T. bisetosa (Coquillett)
and 7. nigricornis (Coquillett) (Goeden
750
Bisse
aeropyles.
Egg of Trupanea signata: pedicel with
1987, 1988; Headrick and Goeden 1991;
Knio et al. 1996a). The single row of aero-
pyles circumscribing the pedicel is similar
to T. nigricornis, whereas, T. bisetosa com-
monly has two rows of aeropyles (Knio et
al. 1996a).
Third instar: White, barrel-shaped, taper-
ing anteriorly, rounded posteriorly; minute
acanthae circumscribe each thoracic and ab-
dominal segment anteriorly, gnathocephalon
conical (Fig. 2A), rugose pads dorsally and
laterally, rugose pads laterad of mouth lu-
men serrated on ventral margin (Fig. 2A-1);
dorsal sensory organ consists of a single
dome-shaped papilla (Fig. 2A-2, B-1); an-
terior sensory lobe bears four sensory or-
gans, lateral sensory organ with a distinct
central papilla (Fig. 2B-2); stomal sense or-
gan ventrolaterad of anterior sensory lobe
(Fig. 2B-3); mouth hooks tridentate; rugose
pads circumscribe prothorax posteriorly of
minute acanthae (Fig. 2C-1), single row of
verruciform sensilla circumscribe prothorax
medially (Fig. 2C-2); anterior thoracic spir-
acles on posterior margin of prothorax, bear
4—5 rounded papillae (Fig. 2C-3, 2D); me-
tathoracic lateral spiracular complex con-
sists of a spiracle and a single verruciform
sensillum; abdominal lateral spiracular
complex consists of a spiracle (Fig. 2E-1),
a verruciform sensillum (Fig. 2E-2), and
placoid-type sensillum (Fig. 2E-3); caudal
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
segment smooth medially, circumscribed
anteriorly by minute acanthae (Fig. 2F-1);
posterior spiracular plates (Fig. 2F-2), with
three ovoid rimae, ca. 0.038 mm in length
(Fig. 2G-1), and four interspiracular pro-
cesses each with 3-6 branches, longest
measuring 0.013 mm (Fig. 2G-2); inter-
mediate sensory complex ventrad of pos-
terior spiracular plates among the minute
acanthae (Fig. 2F-3), consist of a medusoid
sensillum (Fig. 2H-1), and a stelex sensil-
lum (Fig. 2H-2).
Trupanea signata is similar in general
appearance to other described species, i.e.,
Trupanea californica (Headrick and Goe-
den 1991), T. bisetosa and T. nigricornis
(Knio et al. 1996a). The anterior portion of
the prothorax is circumscribed by minute
acanthae and rugose pads, which appear to
be characteristic of the genus Trupanea
(Headrick and Goeden 1991, Knio et al.
1996a). Differences among Trupanea spe-
cies described to date are found in the ab-
dominal lateral spiracular complex. This
complex in T. californica includes a single
verruciform sensillum; in 7. nigricornis,
two verruciform sensilla; and in T. bisetosa,
two verruciform sensilla and a placoid type
sensillum (Headrick and Goeden 1991,
Knio et al. 1996a). Trupanea signata also
differs slightly in the number of branches
in the interspiracular processes; T. califor-
nica and T. bisetosa possess 6—8 branches,
T. nigricornis is similar to T. signata in
having 3—6 branches (Headrick and Goeden
1991, Knio et al. 1996a).
Puparium: Puparium of T. signata shiny
black, elongate-ellipsoidal, anterior end
bears the invagination scar (Fig. 3A-1), and
anterior thoracic spiracles (Fig. 3A-2), cau-
dal segment bears the posterior spiracular
plates (Fig. 3B-1), a band of minute acan-
thae (Fig. 3B-2), and the intermediate sen-
sory complex (Fig. 3B-3). Forty-four pu-
paria of T. signata averaged 3.23 + 0.04
(range, 2.55—3.80) mm in length; 1.48 +
0.03 (range, 1.10—2.35) mm in width.
VOLUME 99, NUMBER 4
1GKU R446
Fig. 2. Third instar of Trupanea signata: (A) gnathocephalon, anterior view, 1—serrated rugose pads, 2—
dorsal sensory organ; (B) anterior sensory lobe, 1—dorsal sensory organ, 2—lateral sensory organ, 3—stomal
sense organ; (C) anterior view, |—prothoracic rugose pads, 2—verruciform sensilla, 3—anterior thoracic spi-
racle; (D) anterior thoracic spiracle; (E) fourth abdominal segment, lateral spiracular complex, 1—spiracle, 2—
verruciform sensillum, 3—placoid-type sensillum; (F) caudal segment, 1—minute acanthae, 2—posterior spi-
racular plate, 3—intermediate sensory complex; (G) posterior spiracular plate, 1—rima, 2—interspiracular pro-
cess; (H) intermediate sensory complex, 1—medusoid sensillum, 2—stelex sensillum.
ee Wier a
1GkKU %240°° ‘Go07° 1paOmmm
Fig. 3. Puparium of Trupanea signata: (A) ante-
rior end, l1—invagination scar, 2—anterior thoracic
spiracles; (B) caudal segment, 1—posterior spiracular
plate, 2—minute acanthae, 3—intermediate sensory
complex.
DISTRIBUTION AND Hosts
The distribution of 7. signata mapped by
Foote et al. (1993) included the western
U.S. north of Mexico and Canada, with this
species recorded from Arizona, California,
Colorado, Nebraska, New Mexico, Oregon,
Texas, and Washington as well as British
Columbia. Foote (1960) and Foote et al.
(1993) reported Anaphalis sp. and Gna-
phalium stramineum Kunth as hosts, be-
sides G. luteo-album reported by Goeden
(1992). Gnaphalium luteo-album is an in-
troduced annual plant species of Old World
origins (Munz 1974). Thus, 7. signata like
T. californica (Headrick and Goeden 1991),
provide examples of indigenous, oligopha-
gous tephritids that have adopted this non-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
indigenous, host-plant species that is con-
generic with native hosts in the southern
California flora (Munz 1974). Besides
Campiglossa genalis (Thomson) on Senecio
spp. (Goeden et al. 1994), few other ex-
amples of such host-plant transfer by indig-
enous, oligophagous, nonfrugivorous Te-
phritidae have been documented (Goeden
1996). The biological significance of the
sweep record for a female of T. signata re-
ported by Goeden (1986) and Foote et al.
(1993) was its location on Santa Cruz Is-
land, not the nonhost, Baccharis pilularis
de Candolle from which it was swept,
which like all-too-many sweep records for
adult, nonfrugivorous Tephritidae are poor
and often misleading indicators of repro-
ductive-host-plant affinities. Accordingly,
T. signata is oligophagous on certain spe-
cies of Anaphalis and Gnaphalium of the
subtribes Cassiniinae and Gnaphaliinae, re-
spectively, and the tribe Gnaphalieae of the
Asteraceae (Bremer 1994). Unlike JT. con-
jJuncta, T. signata apparently is an obligate,
not a facultative gall former, and has not
been reared from flower heads of Anaphalis
or Gnaphalium spp. (Foote 1960, Foote et
al. 1993, Goeden 1983, 1987, 1992, unpub-
lished data). Also, our study sites 1 and 2
were in grazed, disturbed, riparian areas and
site 3 was in a regularly and well irrigated,
roadside flower bed, so all galled plants
were well-watered.
BIOLOGY
Egg.—Eggs were inserted pedicel-last,
singly or side-by-side, in small clusters of
two to four for ca. two-thirds their lengths
in terminal buds of stems or upper axillary
branches (Fig. 4A). Unlike 7. conjuncta,
which also forms galls, the egg clusters
were not glued together posteriorly (Goe-
den 1987).
Larva.—Newly-hatched first instars tun-
nelled basipetally into the pith of the stem
or axillary branch to which they confine
their feeding (Fig. 4B—D). The larvae feed
singly or gregariously in open, elongate
central cavities on proliferating pith paren-
VOLUME 99, NUMBER 4
753
Fig. 4. Life stages of Trupanea signata on Gnaphalium luteo-album: (A) pair of eggs (arrow) inserted in
axillary bud, (B) third instar in feeding cavity below partially dislocated gall cap, (C) exterior view of apical
bud gall, (D) gall of axillary branch with flower head at apex, (E) Four puparia in common feeding cavity below
apical exit channel and gall cap. (F) Female adult at rest. Lines = 1 mm.
chyma (callose tissue), shallowly pitting the
walls, and continually expanding the
lengths and widths of the cavities during all
three stadia (Fig. 4D). The gall cavities re-
mained free of frass and solid waste, except
for the discarded cephalopharyngeal skele-
tons of the first and second instars. The ful-
ly grown third instar extends the cavity dis-
tally by eating out a 5.2 + 0.3 (range, 2-8)
mm_-long, exit tunnel (n = 25) through the
shoot tip to just beneath the apex, leaving
a hollow, tomentum-covered gall “cap,”
which it coats inside with voided liquid fe-
ces. The feces dry, harden, and hold this
cap in place during pupariation (Fig. 4D).
The larva then returns to the main gall cav-
ity, where it and other mature larvae within
pupariate with their heads facing the gall
apex.
Pupa.—One hundred thirteen full-sized
galls collected at all three study sites each
contained an average of 2 + 1 (range, 1—6)
puparia (Fig. 4—E). These galls were elon-
gate-clavoidal in shape and covered with a
754
whitish tomentum (Fig. 4B, C). They av-
eraged 14.9 + 0.6 (range, 6.0—41) mm in
length and 4.8 + 0.1 (range, 1.7—7.8) mm
in widest width near the apices, expanding
gradually distally from a basal, greenish, to-
mentose, side branch or stem, and incor-
porating an average of 3.9 + 0.1 (range, 1—
7) nodes. The galled branch or shoot tips
were not shortened in length, but rather
mainly expanded in width, and like their
ungalled counterparts, bore no or from one
to five vegetative or floral branches laterally
and apically (Fig. 4). Some of these apical
branches were killed when the bud caps
were formed. The cavities within these ma-
ture galls measured 7.1 + 0.3 (range, 3.1—
18.9) mm long by 2.6 + 0.1 (range, 1.3-
6.9) mm in maximum width. Larger galls
typically contained the most puparia, which
lay freely, touching laterally or apically,
within the common cavity (Fig. 4E).
Adult.—Adults emerged through the
same exit tunnel after pushing aside the gall
cap. Adults were long-lived under insectary
conditions, as males averaged 48 + 15
(range, 21—158) days, and 18 females (Fig.
4F) averaged 74 + 11 (range, 31-172)
days. A 7 week-old female contained 35
full-size ova, and eight 3—4 week-old fe-
males laid an average total of 24 + 4
(range, 7-34) eggs in ovipositional cagings.
No free-living adults were observed in na-
ture.
In petri dish arenas, both sexes displayed
synchronous and asynchronous supinations
along with wing vibrations and hamation.
During asynchronous supination, one wing
was extended forward 90° to a point per-
pendicular to the body, and supinated to 90°
with respect to the substrate. As the wing
was extended it was vibrated in a plane par-
allel to the wing blade faster than was ob-
served with other Trupanea spp. studied
(Headrick and Goeden 1994). When the
wing reached its maximum forward posi-
tion, it was held for ca. 1 second, then re-
turned to the resting position, flat upon the
dorsum, and held still while the other wing
was extended. Male courtship displays be-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
gan in the mornings, usually after 0900 h
PST and ended ca. midday. The courtship
display was similar to that described for
other species of Trupanea by Headrick and
Goeden (1991, 1994) and Knio et al.
(1996b). Males of T. signata regularly hung
upside down from the covers of arenas
when displaying and distending their ab-
domens. When the abdomen was distended,
it was held flexed and synchronous or asyn-
chronous wing displays were exhibited;
however, when a female was near, males
always reverted to synchronous wing exten-
sions. Mating was not observed in this spe-
cies, but see Headrick and Goeden (1994)
for descriptions of mating behaviors com-
mon to several southern California species
of Trupanea.
Seasonal history.—The life cycle of T.
signata in southern California follows the
aggregative pattern in which the long-lived
adults in reproductive diapause overwinter
probably in riparian habitats and aggregate
on preblossom host plants during the fol-
lowing spring to mate and reproduce
(Headrick and Goeden 1994). A second
late-summer or early-fall generation repro-
duces on flowering shoots of G. luteo-al-
bum, and another generation or two may be
produced on the above-mentioned, or as yet
unidentified, alternate host plants, especial-
ly at higher elevations.
Natural enemies.—Two species of Hy-
menoptera were reared from puparia of T.
signata as solitary, larval-pupal endopar-
asitoids: Eurytoma sp. (Eurytomidae) and
Pteromalus sp. (Pteromalidae). Among 49
adult parasitoids recovered, nine (18%)
were Eurytoma sp. and 40 (82%) were
Pteromalus sp.
ACKNOWLEDGMENTS
Once again we sincerely thank A. C.
Sanders, Curator of the Herbarium, Depart-
ment of Botany and Plant Sciences, Uni-
versity of California, Riverside, for identi-
fication of plants from southern California
mentioned in this paper. The parasitoids
were identified by Harry E. Andersen, Hun-
VOLUME 99, NUMBER 4
tington Beach, California. We also are
grateful to EK L. Blanc and D. H. Headrick
for their helpful comments on and contri-
butions to earlier drafts of this paper.
LITERATURE CITED
Bremer, K. 1994. Asteraceae Cladistics & Classifica-
tion. Timber Press, Inc. Portland, Oregon.
Cavender, G. L. and R. D. Goeden. 1982. Life history
of Trupanea bisetosa (Diptera: Tephritidae) on
wild sunflower in southern California. Annals of
the Entomological Society of America 75: 400—
406.
Foote, R. H. 1960. A revision of the genus 7rupanea
in America north of Mexico. U.S. Department of
Agriculture Technical Bulletin 1214. 29 pp.
Foote, R. H. and E L. Blanc. 1963. The fruit flies or
Tephritidae of California. Bulletin of the Califor-
nia Insect Survey 7, 117 pp.
Foote, R. H., E L. Blanc, and A. L. Norrbom. 1993.
Handbook of the Fruit Flies (Diptera: Tephritidae)
of America North of Mexico. Cornell University
Press, Ithaca, New York.
Goeden, R. D. 1983. Initial host-plant records for five
species of fruit flies from southern California
(Diptera: Tephritidae). Proceedings of the Ento-
mological Society of Washington 85: 275-281.
1986. New records of Tephritidae (Diptera)
from Santa Cruz Island, California. Pan-Pacific
Entomologist 62: 326-328.
1987. Life history of Trupanea conjuncta
(Adams) on Trixis californica Kellogg in southern
California (Diptera: Tephritidae). Pan-Pacific En-
tomologist 63: 284-291.
1988. Life history of Trupanea imperfecta
(Coquillett) on Bebbia juncea (Bentham) Greene
in the Colorado Desert of southern California
(Diptera: Tephritidae). Pan-Pacific Entomologist
64: 345-351.
. 1992. Analysis of known and new host rec-
ords for Trupanea from California (Diptera: Te-
phritidae). Proceedings of the Entomological So-
ciety of Washington 94: 107-118.
. 1996. Symphagy among florivorous fruit flies
(Diptera: Tephritidae) in southern California. /n
Dettner, K. ed., Vertical Food Web Interactions:
Evolutionary Patterns and Driving Forces. Eco-
logical Studies 109, Springer-Verlag, Heidelberg,
Germany.
Goeden, R. D., D. H. Headrick, and J. A. Teerink.
1994. Life history and description of immature
stages of Paroxyna genalis (Thomson) (Diptera:
Tephritidae) on native Asteraceae in southern Cal-
155
ifornia. Proceedings of the Entomological Society
of Washington 96: 612-629.
Goeden, R. D. and J. A. Teerink. 1996a. Life histories
and descriptions of adults and immature stages of
two cryptic species, Aciurina ferrugenia (Doane)
and A. michaeli new species (Diptera: Tephritidae)
on Chrysothamnus viscidiflorus (Hooker) Nuttall
in southern California. Proceedings of the Ento-
mological Society of Washington 98: 415—438.
. 1996b. Life history and descriptions of adults
and immature stages of Aciurina idahoensis
Steyskal (Diptera: Tephritidae) on Chrysothamnus
viscidiflorus (Hooker) Nuttall in southern Califor-
nia. Proceedings of the Entomological Society of
Washington 98: 681-694.
. 1996c. Life history and descriptions of adults
and immature stages of Aciurina semilucida
(Bates) (Diptera: Tephritidae) on Chrysothamnus
viscidiflorus (Hooker) Nuttall in southern Califor-
nia. Proceedings of the Entomological Society of
Washington 98: 752-766.
1997a. Notes on life histories and descrip-
tions of adults and immature stages of Proceci-
dochares kristineae and P. lisae new spp. (Dip-
tera: Tephritidae) on Ambrosia spp. in southern
California. Proceedings of the Entomological So-
ciety of Washington 99: 67-88.
1997b. Life history and description of im-
mature stages of Procecidochares anthracina
(Doane) (Diptera: Tephritidae) on Solidago cali-
fornica Nuttall in southern California. Proceed-
ings of the Entomological Society of Washington
99: 180-193.
Headrick, D. H. and R. D. Goeden. 1991. Life history
of Trupanea californica Malloch (Diptera: Te-
phritidae) on Gnaphalium spp. in southern Cali-
fornia. Proceedings of the Entomological Society
of Washington 93: 559-570.
1994. Reproductive behavior of California
fruit flies and the classification and evolution of
Tephritidae (Diptera) mating systems. Studia Dip-
terologica 1(2): 194-252.
Hickman, J. C. (ed.) 1993. The Jepson Manual. Uni-
versity of California Press. Berkeley and Los An-
geles.
Knio, K. M., R. D. Goeden and D. H. Headrick.
1996a. Descriptions of immature stages of Tru-
panea nigricornis and T. bisetosa (Diptera: Te-
phritidae) from southern California. Annals of the
Entomological Society of America 89: 1-11.
. 1996b. Comparative biologies of the cryptic,
sympatric species, Trupanea nigricornis and T.
bisetosa (Diptera: Tephritidae) in southern Cali-
fornia. Annals of the Entomological Society of
America 89: 252-260.
Munz, P. A. 1974. A Flora of Southern California.
University of California Press, Berkeley.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, p. 756
NOTE
Celticecis, a Genus of Gall Midges (Diptera: Cecidomyiidae),
Newly Reported for the Western Palearctic Region
Many Holarctic genera of trees and
shrubs are host over much of their ranges
to particular genera of Cecidomyiidae. As
examples, willows host gall midges of Rab-
dophaga and Iteomyia, oaks host Macro-
diplosis and Polystepha, and birches host
Semudobia in both the Nearctic and Pale-
arctic Regions. So many instances of this
pattern are known for gall midges that when
a gall midge genus is recorded from only
part of the range of a Holarctic plant genus,
the partial absence may be suspected to re-
flect a lack of collecting. There are excep-
tions, one being beech. This tree is infested
by gall midges of the genus Mikiola in Eu-
rope and Japan but apparently not in North
America. Mikiola may once have occured
in North America and become extinct due
to the relatively narrow bottleneck beech
passed through during the Pleistocene when
it was restricted to southern North America.
Hackberries, the genus Celtis, are hosts
in North America to at least 10 species of
gall midges of the genus Celticecis that
cause complex leaf and twig galls of some-
times bizarre shape (Gagné 1989, The
Plant-Feeding Gall Midges of North Amer-
ica, Cornell University Press, Ithaca, New
York, xi and 356 pp., 4 pls.). Some of these
species were described 100 years ago, al-
though they were combined into a single
genus only recently (Gagné 1983, Proceed-
ings of the Entomological Society of Wash-
ington 85: 435—438). Celticecis is known
from Japan (Moser 1965, New York State
Museum and Science Service Bulletin 402:
i-iv, 1-95 (as Phytophaga); Yukawa and
Tsuda 1987, Kontya 55: 123-131), which
extends the range of Celticecis into the east-
ern Palearctic. One of us (JCM), pursuing
a long-term interest in hackberry galls, ob-
tained from Prof. K. Browicz of the Insti-
tute of Dendrology, Kornik, Poland, galled
leaves of Celtis tournefortii Lam. collected
in Hisarcik, Kayseri, Turkey. On the upper
surface of the leaf the galls are about 1.5
mm in height and width and consist of an
outer, raised ring and a central, rounded
cone protruding from the the center of the
ring. On the lower surface of the leaf the
gall is a simple convexity about 1.0 mm in
height by 1.5 mm in width. Second instars
of a presumably undescribed species of
Celticecis were extracted from these galls.
This new record extends the natural range
of Celticecis into the western Palearctic.
That Celticecis has not yet been found in
relatively well-collected Europe may mean
that it became extinct there during the Pleis-
tocene, as might have the gall midges on
beech in North America. The galls and the
larvae excised from them are deposited in
the National Museum of Natural History,
Washington, DC.
Raymond J. Gagné, Systematic Entomol-
ogy Laboratory, Retired, PSI, Agricultural
Research Service, USDA, c/o U. S. National
Museum NHB 168, Washington, D. C.
20560, U.S.A., and John C. Moser, Re-
search Entomologist, Emeritus, Southern
Research Station, Forest Insect Research,
2500 Shreveport Hwy., Pineville, LA
FL3S6O, WISA.-
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 757-758
NOTE
Range Expansion of the Red Imported Fire Ant, Solenopsis invicta Buren
(Hymenoptera: Formicidae), into New Mexico and Extreme Western Texas
The red imported fire ant, Solenopsis in-
victa Buren, has greatly expanded its dis-
tribution in the United States since its ac-
cidental introduction into the United States
from South America in the 1930’s (Buren
et al. 1974. Journal of the New York En-
tomological Society 82: 113-124). It cur-
rently occurs throughout the southeastern
United States from southeastern Virginia
(Waller and Vander Meer. 1993. Associa-
tion Southeastern Biologists Bulletin 40:
88), south to Florida (Callcott and Collins.
1996. Florida Entomologist 79: 240-251)
west to Brownsville, Texas (Allen et al.
1993. Southwestern Entomologist 18: 315—
316) and Midland, Lubbock, and Ector
counties, Texas (Cokendolpher and Phillips.
1989. Southwestern Naturalist 34: 443-
449; Porter et al. 1991. Journal of Econom-
ic Entomology 84: 866—874; Callcott and
Collins 1996). Solenopsis invicta is actually
a junior synonym of S. wagneri Santschi
(Bolton. 1995. A New General Catalogue
of the Ants of the World, Harvard Univer-
sity Press, p. 388), but we prefer to us S.
invicta, as the name is so well established.
Introduced fire ants can be separated easily
from our native fire ants by the presence of
a median tooth on the anterior edge of the
clypeus.
We report the presence of the red im-
ported fire ant in New Mexico and El Paso,
Texas. This species will undoubtedly fur-
ther infest urban environments in the south-
western United States and finally invade
southern California, where it will probably
become a major pest.
Specimens were found infesting a home
at Taylor Ranch, a west side suburb of Al-
buquerque, New Mexico, during the sum-
mer of 1994. They were apparently brought
into the house in a large planter that re-
cently arrived from the Gulf Coast. We
know of at least eight cases during 1994
and 1995 when trucks carrying cargo from
the southeastern United States were inspect-
ed and stopped at the New Mexico/Arizona
border. In six cases the trucks were taken
to Albuquerque where they were fumigated.
In the two other cases, the trucks were fu-
migated in Gallup, New Mexico. We know
of no cases of trucks being stopped or fu-
migated during 1996. Discussions with pest
control operators indicate that these eight
cases are only a small fraction of the in-
stances of fires ants found in commerce in
New Mexico. This species was also col-
lected at the ghost town of Steins, New
Mexico (D. Richman, personal communi-
cation). Apparently a backhoe contaminated
with soil was refused entry into Arizona
and returned to the nearby off-ramp where
the soil was removed. Steins is a very arid
site, and it would no be expected that S.
invicta could survive in such a site. The im-
ported fire ant is also found in El] Paso, Tex-
as, the westernmost part of Texas. It occurs
on the campus of the University of Texas
where it nests at the bases of trees. It was
also found near the Rio Grande River in El
Paso in 1989 (R. Worthington, personal
communication). It is not common in El
Paso at the present time, but we expect it
to become more common the this mesic,
urban environment.
The red imported fire ant is not expected
to become a major pest in New Mexico and
western Texas, but these sites will undoubt-
edly serve as “‘stepping stones” for the in-
vasion of the ant into California.
We thank three anonymous reviewers for
helpful comments. Voucher specimens will
be deposited in the National Museum of
Natural History, Smithsonian Institution,
758
Washington, D.C., the collection of Texas
A&M University, College Park, and in our
Laboratory for Environmental Biology.
William P. MacKay, Laboratory for En-
BOOK
Brethren of the Net: American Entomol-
ogy, 1840-1880. By W. Conner Soren-
sen. The University of Alabama Press,
Tuscaloosa, Alabama, U.S.A. 1995. 357
pp., cloth. ISBN 0-8173-0755-9 $59.95.
The idea of insect collectors during mid-
Victorian America may not generate much
initial enthusiasm for some readers. How-
ever, if one is interested in his or her en-
tomological roots or the contributions of
American entomology and entomologists to
science, Brethren of the Net is an essential
read. Sorensen’s book provides a fascinat-
ing glimpse of the events and contributions
that have shaped current scientific thinking
and research not only in entomology but in
other biological sciences as well. The book
contains 12 chapters, two appendices, ab-
breviations, notes, bibliography, and in-
cludes several black and white plates and
photographs.
The first chapter (Entomology in the
American Context) highlights the period be-
fore the 1840’s. Naturalists like Reaumur
and Linnaeus were instrumental in estab-
lishing entomology as a special branch in
natural history with its own nomenclature,
literature, and community of experts. Lin-
naeus’ approach to entomology provided a
system of manageable terms for the disci-
pline. While European entomologists flour-
ished and occupied a niche in European
zoological science, fewer resources were
available for their American counterparts.
By the turn of the 18th century, the situa-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
vironmental Biology, Centennial Museum,
The University of Texas, El Paso, TX
79968, U.S.A. and Richard Fagerlund, De-
partment of Biology, University of New
Mexico, Albuquerque, NM 87131, U.S.A.
PROC. ENTOMOL. SOC. WASH
99(4), 1997, pp. 758-761
REVIEW
tion had begun to change. This was due in
part to the establishment of various insti-
tutions devoted to natural history in the
Philadelphia, Boston—Cambridge, and to a
lesser degree New York area. The careers
of six Americans (John Abbot, Thaddeus
William Harris, John Eatton LeConte, Fred-
erick Valentine Melsheimer, John E Mels-
heimer, William D. Peck, and Thomas Say)
who figured prominently before the gener-
ation of 1840 are emphasized. These early
workers were among the first Americans to
specialize to a similar degree as the Euro-
pean entomologists. American entomolo-
gists were no longer content simply to col-
lect specimens for the Europeans. American
entomologists were coming in closer con-
tact with one another but they lacked estab-
lished institutions for practicing scientific
entomology.
Chapter Two concentrates on the first
American entomological society devoted to
the study of insects—The Entomological
Society of Pennsylvania. Here the emphasis
is on the members that formed the nucleus
of the group. Though there were no formal
requirements for membership to the orga-
nization, the society was somewhat exclu-
sive. The society’s correspondence with
other individuals with an interest in ento-
mology was very significant. It served as a
vital transition institution that bridged a
time between a few isolated investigators
and the time of specialists associated with
large scientific institutions. Members of
The Entomological Society of Pennsylvania
VOLUME 99, NUMBER 4
aspired to advance American entomology in
insect systematics primarily by having
Americans publish descriptions of Ameri-
can insects. This desire was probably best
reflected in John LeConte’s announcement
of the “‘American Entomological Declara-
tion of Independence.’’ Eventually Le-
Conte’s declaration materialized with the
publication of Melsheimer’s catalog of Co-
leoptera. However, with publication of the
catalog, the Society began to fade. The So-
ciety’s main objective in establishing pri-
ority for American names in Melsheimer’s
catalog had been secured.
Of Cabinets and Collections is the title
of the third chapter. This chapter is devoted
to the details involved in the establishment
of the major insect collections of the mid-
19th century. The importance of the ex-
panding entomological collections housed
at the Museum of Comparative Zoology at
Harvard University, the Academy of Natu-
ral Sciences of Philadelphia, and the Smith-
sonian Institution are highlighted. A grow-
ing entomological community along with
the establishment of these collections was
substantial. The westward expansion of the
American republic resulted in numerous
collection expeditions that yielded large
numbers of specimens. A modicum of stan-
dardization for insect cabinets and pins and
the widespread use of preservatives within
the collections kept pests from destroying
the holdings. The financial support from
state and federal governments and wealthy
patrons were important to the entomologi-
cal collections, as was the donation of large
private collections. This was especially the
case for the Smithsonian’s entomological
holdings when C. V. Riley donated his col-
lection (estimated at the time to be the larg-
est general collection in the country) in
1885. By this time, Americans were well
on their way to establishing entomological
institutions that were comparable to those
enjoyed by their European counterparts.
Agricultural Entomologists and Institu-
tions (Chapter 4) elaborates on the trans-
formation of those early “‘amateur’’ ento-
Way
mological investigators to paid profession-
als with an emphasis on public service and
practical application. Funding for agricul-
tural entomology and the establishment of
instruction of the subject at various insti-
tutions was important during this time. In
1863, a course in agricultural entomology
taught by Manley Miles at Michigan Agri-
cultural College was apparently the first of
its kind. By the early 1870’s, course offer-
ings in agricultural entomology appeared in
half a dozen colleges across the United
States. The fact that the level of support in
the United States rose from the great agri-
cultural changes in the nation is notewor-
thy. The states, federal government, and ag-
ricultural colleges offered multiple institu-
tional possibilities for those seeking such
support. This support translated into sub-
stantial expansion of employment for en-
tomologists.
Chapters 5—7 pertain to the topic of the
‘“‘balance of nature’? or “‘economy of na-
ture’? theory and specific insects that
shaped American economic entomology.
The balance of nature theory presupposed
that God regulated the world through nat-
ural checks and balances; it further assumed
a harmonious relationship among all living
species. Such a species has traits that “‘en-
able it to maintain its essential shape (mor-
phology) and to inhabit its geographic
range despite challenges form the other spe-
cies or from the environment.”’ The balance
was “‘upset’’ by either the introduction of
nonnative species or crop monoculture.
Chapters six and seven cover in detail spe-
cific insects (plum curculio, the Hessian fly,
the Colorado potato beetle, and the Rocky
Mountain locust (Chapter 7)) that interrupt-
ed this “‘balance”’ and led to significant ad-
vances for the entomological community.
During 1840—1870 there was much interest
in the relationship between insects and
crops, the destructive outbreaks of insect
pests, and various reasons for upsets in the
balance. State and agricultural entomolo-
gists were called upon to prove or dispel
information presented in the popular agri-
760
culture press at the time. The entomological
reports usually included aspects of insect
bionomics along with illustrations followed
by suggestions for control measures.
Through this work, entomologists deter-
mined which insect species was “‘friend”’ or
‘foe.’ Also, during this period was a great
debate on the “bird question”’ or the rela-
tionship between insects, birds, and crops.
Along with ornithologists, entomologists
were called upon to determine precise feed-
ing habits of various birds. Most farmers
and agriculturists learned to rely on the
work of entomologists to provide reliable
information on various insect life cycles.
Entomologists were then expected to sug-
gest strategies to manipulate the balance of
nature for the benefit of farmers.
Profile of the American Entomological
Community About 1870 (Chapter 8) is an
extremely detailed examination of the de-
mographics of entomologists in the later
19th century. The chapter provides quanti-
tative data on the education background,
place of birth, attended institutions, etc. for
entomologists at that time. Interesting is a
section on the active role of several pio-
neering women in entomology (e.g., Sara
McBride, Maria Mitchell, Mary Murtfelt,
Emily Smith, Charlotte Taylor, and Mary
Treat) during a time when the discipline
was male dominated. The entomological
community of the 1870’s had grown to
about 900 serious investigators of which
approximately 100 functioned as a publish-
ing elite.
Chapter 9 (Acceptance and Implications
of Evolution) discusses the important role
American entomologists played in the sup-
port of evolutionary theory. This chapter
highlights the various debates and contro-
versies within the entomological commu-
nity then. Outspoken advocates such as
Walsh and Riley helped promote evolution-
ary theory, but the fact that many American
entomologists were field-oriented allowed
them to observe the great geographic and
climatic variety of the North American con-
tinent. This coupled with an emphasis of
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
agricultural entomology that many Ameri-
can workers shared, reinforced their under-
standing of the biological and ecological as-
pects of insects. What is particularly note-
worthy is how quickly and with near una-
nimity, American entomologists accepted
evolutionary theory (Darwinian or “‘Neo-
Lamarckians’’) and incorporated it in their
work. The Darwinian revolution placed en-
tomologists in the center of the new field
of evolutionary biology.
Chapters 10 and 11 are examples of two
prominent evolutionary biology studies by
American entomologists in the 19th centu-
ry. Chapter 10 concentrates on the search
for evolutionary explanations of variation
and polymorphism of Lepidoptera (William
Henry Bates and Polymorphism in Butter-
flies) while chapter 11 (The Yucca Moth)
focuses on the coevoution of plants and an-
imals. Both chapters underscore the contri-
butions of American entomologists toward
acceptance of evolutionary theory in the
scientific community.
The Debate over Entomological Nomen-
clature (Chapter 12) reflects the nomencla-
tural crisis that faced the entomological
community in the 1870’s. New Darwinian
principles as applied to the definition of
species and their systematic arrangement
brought the issue to the foreground. The
radical “‘splitting’’ of genera by some en-
tomologists facilitated the debate. For en-
tomology at the time, the inadequacy of no-
menclatural rules presented a problem. It
was argued by some that entomologists
faced more pressing needs than other zool-
ogists in the clarification of the rules of no-
menclature. Debates such as these moved
American entomologists to a prominence
and special point of influence in interna-
tionally recognized rules of nomenclature.
The final chapter (Conclusion) summa-
rizes the progress that American entomol-
ogy made in such a short time during the
1800’s. By 1880, entomologists were the
largest single group of zoologists in North
America. Along with the number of ento-
mologists, knowledge of the North Ameri-
VOLUME 99, NUMBER 4
can insect fauna had also grown apprecia-
bly—not only in the total described species
but also in the number of life history stud-
ies. The number of institutions teaching en-
tomology or associated with the science too
had grown. Entomological collections and
outlets for publications had expanded sub-
stantially during the period. A discipline
that began with a few amateur naturalists
had developed into a fully professional
group.
Brethren of the Net will have a special
appeal for the student of history, science, or
both and would make a wonderful compan-
ion to such books as Mallis’ American En-
tomologists (Rutgers University Press,
1971) and Evan’s Pioneer Naturalists (Hen-
ry Holt and Co., Inc., 1993). The work is
well written, researched, amply footnoted,
and I heartily recommend it. Sorensen has
elaborated on events and individuals that
761
have not only had an important influence
on entomology but on science overall. Al-
though some of these people or incidents
are often forgotten, they should not be. Sor-
ensen has gone a long way to remedy this.
He has resurrected those people and events,
and given the reader tremendous insight
into the circumstances surrounding impor-
tant episodes in science’s past. As scientists
we often forget that science too builds upon
the work of others. Scientific knowledge
does not spontaneously generate. Brethren
of the Net goes a long way in proving this
as it underscores the influence of previous
events on our current work.
I thank Andrew Jensen and Melissa Mil-
ler for comments on an early draft of this
review.
Gary L. Miller, 338 Laurel Ave., Laurel,
MD 20707 U.S.A.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 762-771
OBITUARY
Jenaro Maldonado Capriles
1919-1995:
Biographical Sketch, Patronyms in His Honor, and List of Publications
Jenaro Maldonado Capriles (Fig. 1), 76,
died on 22 November 1995, in San Anto-
nio, Texas. He is survived by his wife, Car-
men; their children, Ricardo and Maria Te-
resa; a grandaughter, Jacqueline Marisa;
and by many friends and colleagues.
Jenaro, as most of his colleagues called
him, was born in Yauco, Puerto Rico, on 17
January 1919, where he attended primary
and secondary school. In 1939, Maldonado
finished his Bachelor’s Degree in Agricul-
tural Sciences at the “‘Colegio de Agricul-
tura y Artes Mecanicas’” (CAAM, now
known as the Recinto Universitario de Ma-
yagtiez, RUM, of the University of Puerto
Rico). He completed a Master’s in Sanitary
Sciences at the “Escuela de Medicina”
(School of Tropical Medicine, now part of
the Escuela de Medicina,’’ UPR) in 1942.
He related to one of us (JSB) that he be-
came interested in medical entomology so
he could work on something that would re-
ally make a difference in people’s health.
He served as an Entomologist for the Bu-
reau of Malaria Control of the ‘‘Departa-
mento Estatal de Salubridad’”’ (State Health
Department) from 1941 to 1948, as part of
efforts to eradicate malaria in Puerto Rico.
Those years yielded his first publication:
‘The fleas of Puerto Rico”’ (1945). In 1948,
he joined the faculty of CAAM as an As-
sistant Professor. Maldonado received his
doctorate from Ohio State University, Co-
lumbus, in 1956. His dissertation was en-
titled “‘A study of some Neotropical leaf-
hoppers (Homoptera: Cicadellidae: Idioce-
rinae)’’; Carl E. Vernard was his Major Pro-
fessor. He viewed his years at OSU as
interesting and enjoyable but all too short.
Dr. Maldonado served as Chairperson of
the Biology Department (CAAM) from
1960—1966 and as a faculty member of the
Ponce School of Medicine from 1977—
1982, where he was Chairman of the De-
partment of Anatomy. He held numerous
additional positions, including academic,
administrative, community, and others, in
Puerto Rico and worldwide.
Maldonado was also an avid field ento-
mologist, frequently going on collecting ex-
peditions to southeast Asia, the Caribbean,
North Africa, and South Africa, among oth-
ers. He was enthusiastic in initiating new
and biologically interesting research, partic-
ularly in the Hemiptera. His taxonomic
work mainly involved the hemipteran fam-
ilies Cicadellidae (Auchenorrhyncha), Mi-
ridae (Heteroptera), and Reduviidae (Het-
eroptera). Like most systematists, he had a
few major projects and many side investi-
gations going on simultaneously, always
pursuing his science with excitement and
enthusiasm, just as traditional country Puer-
to Rican children might open their presents
on Three Kings Day. Even at an advanced
age, he eagerly talked about learning and
applying new technologies to his research,
including scanning electron microscopy,
which he incorporated into some of his later
publications. His 704-page ‘‘Systematic
Catalogue of the Reduviidae of the World”
(1990), undoubtedly, is his major contri-
bution to entomology. Another important
work is his Miridae of Puerto Rico, pub-
lished in 1969 (and supplemented by addi-
tions and corrections in 1991).
Maldonado’s reputation as a concerned
pedagogue and strict teacher always fol-
VOLUME 99, NUMBER 4 763
Fig. 1. Dr. Jenaro Maldonado Capriles at the Department of Entomology, National Museum of Natural
History, Smithsonian Institution, Washington, D.C., 1981.
764
Fig. 2.
Dr. José Ramos at a meeting of the Sociedad Ento-
mologica de Puerto Rico, circa 1980.
Dr. Jenaro Maldonado Capriles (right) with
lowed him. He rewarded students for hard
and careful work and had a profound dislike
for those who were disinterested. He held
similar views about local politicians and
physicians. Along with his wife Carmen, he
was in charge of the Honor’s Program at
CAAM for several years. From the late
1950s, his efforts to further Puerto Rican
education emphasized nursing, medical
technology, numerous biology courses, and
university libraries, mostly at CAAM.
Professor Maldonado was open and en-
thusiastic, and was still vigorously pursuing
his entomological research until the time of
his death. Along with recently deceased
Professor José Ramos (Fig. 2), Maldona-
do’s death constitutes a major loss to En-
tomology in Puerto Rico and the world.
Maldonado’s private insect collection of
approximately 300 Schmidt boxes, involv-
ing more than 18,000 specimens, and most
of his primary types are now housed in the
National Museum of Natural History
Smithsonian Institution (Washington, D.C.).
For a limited time, reprints of Maldona-
do’s more recent papers, except the Cata-
logue, can be obtained by writing to Dr.
Carmen Acosta Maldonado (Calle 6-I-1,
Urbanizacion Aponte, Cayey, PR 00736).
His catalogue can be purchased from the
Caribbean Journal of Science, Special Pub-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
lications, Recinto Universitario de Maya-
gliez, Mayagiiez, PR 00680, at a cost of
$US15.00, plus shipping and handling.
ACKNOWLEDGMENTS
We thank Dr. Carmen Acosta Maldonado
(San Juan Bautista School of Medicine, San
Juan, Puerto Rico) for providing us with a
copy of Jenaro’s Curriculum Vitae. Luis Fe-
lipe Martorell, Silverio Medina-Gaud, and
Mario Pérez-Escolar (Crop Protection De-
partment, University of Puerto Rico, Ma-
yagtiez) provided some details of Professor
Maldonado’s early years. Dr. David L. Den-
linger (Chairman, Department of Entomol-
ogy, Ohio State University, Columbus) also
provided data concerning Maldonado’s
years at OSU. Jo-Anne Hayes (Librarian,
Agricultural Experiment Station, University
of Puerto Rico), Andrew L. Hicks (Boulder,
Colorado), and EK C. Thompson (Systematic
Entomology Laboratory [SEL], ARS,
USDA, c/o National Museum of Natural
History, Washington, D.C.) kindly provided
some of the literature to complete the list
of patronyms. Drs. R. C. Froeschner (Na-
tional Museum of Natural History
[NMNH], Washington, D.C.), D. A. Pol-
hemus (NMNH), D. R. Smith (SEL), and EF
C. Thompson (SEL) read the manuscript
and offered suggestions for its improve-
ment.
PATRONYMS NAMED IN HONOR OF JENARO
MALDONADO-CAPRILES
Alienates maldonadoi Wygodzinsky and
Schmidt. 1991. Bull. Am. Mus. Nat.
Hist. 200: 228. [Heteroptera: Enicoce-
phalidae]. Distribution: Jamaica.
Ambrysus maldonadus [sic] La Rivers.
1954. Entomol. News 65: 197. [Heter-
optera: Naucoridae]. Distribution: Vene-
zuela.
Antias maldonadoi Carvalho. 1982. Rev.
Brasil. Biol. 42(3): 331. [Heteroptera:
Miridae]. Distribution: Dominican Re-
public.
Arenasella maldonadoi Caldwell. 1951. J.
Agric. Univ. Puerto Rico 34: 224.
VOLUME 99, NUMBER 4
[Auchenorrhyncha: Flatidae]. Distribu-
tion: Puerto Rico. [The original spelling,
A. maldonadi, is incorrect and is here
emended (second ‘‘o”’ of patronym add-
ed) a]:
Balocha maldonadoi Kameswara Rao and
Ramakrishnan. 1979. J. Bombay Nat.
Hist. Soc. 76: [Auchenorrhyncha: Cica-
dellidae]. Distribution: India. [The orig-
inal spelling B. maldanadoi is incorrect
is here emended (second “‘a”’ of patron-
ym changed to “‘o’’)].
Coronigoniella maldonadoi Young. 1977.
Tech. Bull. 239, North Carolina Agric.
Exp. Stn. (Raleigh). p. 1020. [Auchen-
orrhyncha: Cicadellidae]. Distribution:
Venezuela.
Deltocoelidia maldonadoi Kramer. 1961.
Proc. Biol. Soc. Wash. 74: 238. [Auche-
norrhyncha: Cicadellidae]. Distribution:
Venezuela.
Diphleps maldonadoi Henry. 1977. Florida
Entomol. 60: 206. [Heteroptera: Miri-
dae]. Distribution: Puerto Rico.
Ectomocoris maldonadoi Cai and Lu. 1991.
Entomotaxonomia 13: 243. [Heteroptera:
Reduviidae]. Distribution: China.
Guanabarea maldonadoi Carvalho. 1984.
Rev. Brasil. Biol. 44(3): 320. [Heterop-
tera: Miridae]. Distribution: Puerto Rico.
Hadria maldonadoi Young. 1977. Tech.
Bull. 239, North Carolina Agric. Exp.
Stn. (Raleigh). p. 974. [Auchenorrhyn-
cha: Cicadellidae]. Distribution: Domin-
ican Republic.
Hermaeophaga maldonadoi Blake. 1965.
Proc. Entomol. Soc. Wash. 67: 109. [Co-
leoptera: Chrysomelidae]. Distribution:
St. Thomas.
Hexatoma maldonadoi Alexander. 1952. J.
New York Entomol. Soc. 60: 245. [Dip-
tera: Tipulidae]. Distribution: Venezuela.
Idioscopus capriliana Viraktamath and
Murphy. 1980. [Auchenorrhyncha: Ci-
cadellidae]. J. Entomol. Res. 4:85. Dis-
tribution: India.
Idioscopus caprilei Webb. 1976. J. Ento-
mol. Soc. S. Afr. 39: 304. [Auchenor-
rhyncha: Cicadellidae]. Distribution: Ni-
765
geria. [This species is now in the genus
Pretioscopus)}.
Lepidocyrtus maldonadoi Mari Mutt. 1986.
Carib. J. Sci. 22 (1—2): 13. [Collembola:
Entomobryidae]. Distribution: Puerto
Rico.
Longurio maldonadoi Alexander. 1953. J.
New York Entomol. Soc. 61: 147. [Dip-
tera: Tipulidae]. Distribution: Venezuela.
Maldonadora Webb. 1983. Bull. Brit. Mus.
Nat. Hist. 47: 228. [Auchenorryncha: Ci-
cadellidae].
Minasmiris maldonadoi Carvalho. 1985.
Rev. Brasil. Zool. 3(4): 163. [Heterop-
tera: Miridae]. Distribution: Colombia.
Monobelus maldonadoi Ramos. 1957. J.
Agric. Univ. Puerto Rico 41: 104. [Au-
chenorrhyncha: Membracidae]. Distri-
bution: Puerto Rico.
Prepops maldonadoi Carvalho and Fontes.
1973. Rev. Brasil. Biol. 33: 543. [Het-
eroptera: Miridae]. Distribution: Domin-
ican Republic.
Sthenaridea maldonadoi Schuh and
Schwartz. 1988. Bull. Am. Mus. Nat.
Hist. 187: 189. [Heteroptera: Miridae].
Distribution: Mexico; widespread in
Neotropics.
Wiloatma caprilesi Webb. 1983. Aust. J.
Zool. Suppl. Series 92: 76. [Auchenor-
rhyncha: Cicadellidae]. Distribution:
Australia.
PUBLICATION LIST
The following is a list of Maldonado’s
157 scientific publications. Most of his pa-
pers deal with various groups of Heterop-
tera, particularly the Reduviidae and Miri-
dae. He also conducted research in other ar-
thropod groups, many of economic impor-
tance: Acari (Ixodida), Hemiptera:
Auchenorrhyncha (Cicadellidae, Issidae,
and Tropiduchidae), Anoplura, Mallophaga,
Coleoptera (Scarabaeidae and Lymexyloni-
dae), and Diptera (Culicidae, Syrphidae,
and Tabanidae). In addition, Maldonado
wrote about a dozen popular articles that
are not listed. We also note that there are
several other papers in preparation that will
766
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
include Maldonado as an author, but these
are
not far enough along to be included
here.
. Maldonado Capriles, J.
. Maldonado Capriles, J. 1945. The fleas of Puerto
Rico. Puerto Rico J. Public Health Trop. Med.
21:173—183 [also in Spanish 21: 184-192].
. Maldonado Capriles, J. 1948. Three new neo-
tropical species of the reduviid genus Ploiaria
Scopoli (Hemiptera). Proc. Entomol. Soc. Wash.
50: 18-22.
1948. A new Enico-
cephalus from Puerto Rico. Proc. Entomol. Soc.
Wash. 50: 159-160.
. Rehn, J. W. H., J. Maldonado Capriles, and J. M.
Henderson. 1950. Field studies on the bionomics
of Anopheles albimanus. Parts I and III: Diurnal
resting places—Progress Report. J. Natl. Malaria
Soc. 9: 268-279.
. Drake, C. J. and J. Maldonado Capriles. 1952.
Water striders from Territorio Amazonas, Vene-
zuela. Great Basin Nat. 12: 47—54.
. Fox, I. and J. Maldonado Capriles. 1953. Light
trap studies on mosquitoes and Culicoides in
western Puerto Rico. Mosquito News 13: 165—
166.
. Maldonado Capriles, J. 1953. Five new neotrop-
ical species of Ghilianella (Hemiptera, Reduvi-
idae). Proc. Entomol. Soc. Wash. 55: 189-195.
. Maldonado Capriles, J. 1953. Redescription of
the genus Burtinus Stal and description of a new
species from Puerto Rico (Hemiptera: Coreidae).
Proc. Entomol. Soc. Wash. 55: 40—44.
. Drake, C. J. and J. Maldonado Capriles. 1954.
Puerto Rican water-striders (Hemiptera). Proc.
Biol. Soc. Wash. 67: 219-221.
. Maldonado Capriles, J. 1954. A note on the ge-
nus /diotettix Osborn (Homoptera, Cicadellidae).
Proc. Entomol. Soc. Wash. 56: 247-250.
. Maldonado Capriles, J. 1955. Cryptotylus stonei,
a new tabanid from Venezuela (Diptera, Tabani-
dae). Proc. Entomol. Soc. Wash. 57: 189-190.
. Maldonado Capriles, J. 1955. Four new Vene-
zuelan reduviid bugs. Proc. U.S. Natl. Mus. 104:
105-113.
. Drake, C. J. and J. Maldonado Capriles. 1955.
Some pleids and water-striders from the Domin-
ican Republic (Hemiptera). Bull. Brooklyn En-
tomol. Soc. 51: 53-56.
. Drake, C. J. and J. Maldonado Capriles. 1955.
New apterous Aradidae from Puerto Rico (He-
miptera). J. Wash. Acad. Sci. 45:289-294.
. Drake, C. J. and J. Maldonado Capriles. 1956. A
remarkable new Rhagovelia from the Dominican
Republic (Hemiptera: Veliidae). Bull. S. Cal.
Acad. Sci. 55: 14-17.
. Maldonado Capriles, J. W. E Pippin, and M. L.
Kuns. 1958. An annotated check list of the mos-
Wie
18.
i)
iw)
23)
N
nn
26.
2k:
28.
2).
30.
quitoes of Mona Island, Puerto Rico, and the lar-
va and male of Aedes obturbator D. & K. (Dip-
tera, Culicidae). Proc. Entomol. Soc. Wash. 60:
65-68.
Maldonado Capriles, J. and S. M. Rafi. 1958. Re-
sults of malariometric surveys performed be-
tween 1939 & 1957 in northern West Pakistan.
Pakistan J. Health 8: 87-90.
Drake, C. J. and J. Maldonado Capriles. 1959. A
new tingid from Pakistan (Hemiptera). Bull.
Brooklyn Entomol. Soc. 54: 25-26.
. Mariota Trias, E and J. Maldonado Capriles.
1959. Insectos perjudiciales a la cafia de azticar
y su combate. Rev. Agric. Puerto Rico 46: 67—
74.
. Maldonado Capriles, J. and A. S. Nasir. 1960.
DDT-resistant adults of Anopheles subpictus in
the Lahore District of West Pakistan. Mosquito
News 20: 52-54.
. Maldonado Capriles, J. 1960. Assassin bugs of
the genus Ghilianella in the Americas (Hemip-
tera, Reduviidae, Emesinae). Proc. U.S. Natl.
Mus. 112: 393—450.
. Maldonado Capriles, J. 1961. Studies on Idiocer-
inae leafhoppers: I./diocerinus Baker, 1915, syn-
onym of Balocha Distant, 1908, and notes on the
species of Balocha (Homoptera: Cicadellidae).
Proc. Entomol. Soc. Wash. 63: 300-308.
Maldonado Capriles, J. and C. A. Navarro. 1962.
Number of insect species reported from Puerto
Rico. Carib. J. Sci. 2: 22—26.
. Gurney A. B. and J. Maldonado Capriles. 1962.
On the cricket Hygronemobius alleni (Morse),
with special reference to a first record of the ge-
nus in Puerto Rico (Orthoptera, Gryllidae). Car-
ib. J. Sci. 2: 156-158.
. Maldonado Capriles, J. and T. H. Farr. 1962. On
some Jamaican Triatominae and Emesinae (Re-
duviidae: Hemiptera). Proc. Entomol. Soc. Wash.
64: 187-194.
Rivero, J. A., J. Maldonado Capriles, and H.
Mayorga. 1963. On the habits and food of Eleu-
therodactylus karlschmidti Grant. Carib. J. Sci.
3: 25-27.
Maldonado Capriles, J. 1963. A new species of
Ghilianella from Haiti (Hemiptera Reduviidae).
Proc. Entomol. Soc. Wash. 65: 64—66.
Maldonado Capriles, J. 1964. Studies on Idiocer-
inae leafhoppers: II]. The Indian and Philippine
species of /diocerus and the genus /dioscopus
(Homoptera: Cicadellidae). Proc. Entomol. Soc.
Wash. 66: 89-100.
Maldonado Capriles, J. 1965. Studies on Idiocer-
inae leafhoppers: III. On Singh-Pruthi’s Indian
species of diocerus (Homoptera, Cicadellidae).
Proc. Entomol. Soc. Wash. 67: 244-246.
Drake, C. J. and J. Maldonado Capriles. 1965. A
VOLUME 99, NUMBER 4
Sill
52)
33)
35:
36.
Sie
38.
39)
40.
41.
42.
43.
44.
45.
new lacebug from Haiti (Hemiptera: Tingidae).
J. Kansas Entomol. Soc. 38: 317-319.
Maldonado Capriles, J. 1966. Sobre algunos
emesinos (Reduviidae, Hemiptera) venezolanos y
descripcié6n de nuevas especies. Mem. Soc.
Cienc. Natur. La Salle 26: 195-204.
Maldonado Capriles, J. 1966. A note on Zan-
chisme (Hemiptera: Miridae). Proc. Entomol.
Soc. Wash. 68: 21-24.
Maldonado Capriles, J. and P. H. van Doesburg,
Jr. 1966. On some Emesinae from Dutch Guiana
(Surinam), with a new species (Reduviidae).
Proc. Entomol. Soc. Wash. 68: 325-329.
. Maldonado Capriles, J. and C. A. Navarro. 1967.
Additions and corrections to Wolcott’s “Insects
of Puerto Rico.”’ Carib. J. Sci. 7: 45-64.
Maldonado Capriles, J. 1968. Nuevos y viejos
emesinos neotropicales (Hemiptera, Reduviidae).
Mem. Soc. Cienc. Nat. La Salle 28: 265-280.
Maldonado Capriles, J. 1968. A new genus and
species of Deltocephalinae from Puerto Rico
(Homoptera: Cicadellidae). Proc. Entomol. Soc.
Wash. 70: 35-37.
Maldonado Capriles, J. 1968. Studies in Idiocer-
inae leafhoppers: IV. A new species of Balocha
and one of Pedioscopus, mimics. Proc. Entomol.
Soc. Wash. 70: 97—100.
Maldonado Capriles, J. 1969. The Miridae of
Puerto Rico (Hemiptera). Tech. Paper No. 45.
Univ. Puerto Rico Agric. Exp. Stn. 133 pp.
Maldonado Capriles, J. 1970. Descriptions of
new species of the genus Termatophylidea with
a key to the known species (Hemiptera: Miridae).
Proc. Entomol. Soc. Wash. 72: 119-126.
Maldonado Capriles, J. 1970. Studies on Idiocer-
inae leafhoppers: VI, New species of Balocha
from the Papuan Subregion (Homoptera: Cica-
dellidae). Pacific Insects 12: 297-302.
Maldonado Capriles, J. 1970. New species in the
genus Sericophanes Reuter (Hemiptera: Miri-
dae). Proc. Entomol. Soc. Wash. 72: 98—106.
Maldonado Capriles, J. 1971. Studies on Idiocer-
inae leafhoppers: VII: Concerning the Ethiopian
genus Rotifunkia China 1926 (Hom., Cicadelli-
dae). Ann. Entomol. Fenn. 37: 202-204.
Keirans, J. E., C. M. Clifford, and J. Maldonado
Capriles. 1971. Argas (Argas) dulus, new species
(Ixodoidea: Argasidae), from nests of the palm
chat Dulus dominicus in the Dominican Repub-
lic. Ann. Entomol. Soc. Am. 64: 1410-1413.
Maldonado Capriles, J. and S. Medina Gaud.
1971. A simple way to determine the presence of
the stable fly, Stomoxys calcitrans (L.) (Diptera:
Muscidae), on dairy farms in Puerto Rico. J.
Agric. Univ. Puerto Rico 55: 259-260.
Maldonado Capriles, J. and S. Medina Gaud.
1971. Distribution and abundance of the cattle
tail louse, Haematopinus quadripertusus Fahr.
46.
AT.
48.
49.
50.
Site
54.
55.
56.
Si:
58.
767
(Anoplura, Haematopinidae) in Puerto Rico. J.
Agric. Univ. Puerto Rico 55: 516-517.
Maldonado Capriles, J. 1971. Platybasicornis ra-
mosi, a new neotropical genus and species (He-
miptera: Miridae: Hyaliodini). Proc. Entomol.
Soc. Wash. 73: 142-145.
Maldonado Capriles, J. 1971. About Idiocerinae
leafhoppers: V. Balcanocerus, a new genus for
Chunrocerus balcanicus Zakhvatkin, 1946 (Ho-
moptera: Cicadellidae). Proc. Entomol. Soc.
Wash. 73: 184-187.
Maldonado Capriles, J. and P. S. Fiuza Ferreira.
1971. Carvalhomiris brachypterus, a new mirid
genus and species from Colombia (Hemiptera,
Miridae). Rev. Brasil. Biol. 31: 345-347.
Wygodzinsky, P. and J. Maldonado Capriles.
1972. Description of the first genus of physod-
erine assassin bugs (Reduviidae, Hemiptera)
from the New World. Am. Mus. Novitates 2504:
1-7.
Maldonado Capriles, J. 1972. Neotropical Re-
duviidae (Heteroptera) in the Museum of Zool-
ogy of the University of Helsinki, Finland, with
description of new species. Not. Entomol. 52:
47-56.
Maldonado Capriles, J. 1972. Studies on Idiocer-
inae leafhoppers: VIII. The Papuan genus Pe-
dioscopus and two allied new genera from the
Philippine Islands (Homoptera: Cicadellidae).
Pacific Insects 14: 529-551.
. Maldonado Capriles, J. 1972. Studies on idiocer-
ine leafhoppers, IX. Three new genera from the
eastern Oriental Region (Homoptera: Cicadelli-
dae). Pacific Insects 14: 627-633.
. Maldonado Capriles, J. and M. Colon Ferrer.
1973. Severe attack on sugar cane and grasses by
Mocis latipes (Guenée) (Lepidoptera: Noctuidae)
in southwest Puerto Rico. J. Agric. Univ. Puerto
Rico 57: 348-349.
Maldonado Capriles, J. 1973. Studies on Idiocer-
inae leafhoppers: X. Idioscopus nitidulus (Walk-
er), new combination (Homoptera: Cicadellidae).
Proc. Entomol. Soc. Wash. 75: 179-181.
Maldonado Capriles, J. 1973. Parapycnoderes, a
new genus for Pycnoderes porrectus (Distant)
(Hemiptera: Miridae). Proc. Entomol. Soc. Wash.
75: 314-317.
Carvalho, J. C. M. and J. Maldonado Capriles.
1973. Mirideos neotropicais, CL: Descrigao de
trés espécies novas (Hemiptera). Rev. Brasil.
Biol. 33: 39-42.
Maldonado Capriles, J. 1974. Studies on Idiocer-
inae leafhoppers, XI. The neotropical genus
Chunroides (Cicadellidae). Entomol. Mon. Mag.
110: 233-236.
Maldonado Capriles, J. 1974. Dos nuevos redu-
viidos venezolanos (Insecta: Hemiptera: Redu-
768
33).
60.
61.
62.
64.
65.
66.
67.
68.
69.
70.
Wile
2.
V3.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
viidae). Mem. Soc. Cienc. Nat. La Salle 34: 53—
DNS}
Maldonado Capriles, J. 1976. Studies on Idiocer-
inae leafhoppers: XIII: /dioceroides Matsumura
a new genus from Taiwan
(Agallinae: Idiocerinae). Pacific Insects 17: 139—
143.
Maldonado Capriles, J. 1974. The neotropical ge-
nus Melanopleurus (Hemiptera: Lygaeidae).
Proc. Entomol. Soc. Wash. 76: 22-30.
Maldonado Capriles, J. 1974. Studies on Idiocer-
inae leafhoppers XII. /dioscopus clavosignatus
spec. nov. (Homoptera, Cicadellidae). Zool.
Meded. 48: 163-167.
Maldonado Capriles, J. 1975. Studies on Idiocer-
inae leafhoppers, XVI: Pachymetopius Matsu-
mura transferred to Coelidiinae (Homoptera: Ci-
cadellidae). Proc. Entomol. Soc. Wash. 77: 306—
307.
and Anidiocerus,
. Maldonado Capriles, J. 1975. Studies on Idiocer-
inae leafhoppers: XVII: Three new neotropical
genera (Homoptera: Cicadellidae). Proc. Ento-
mol. Soc. Wash. 77: 317-325.
Maldonado Capriles, J. 1975. Two new neotrop-
ical Stenolemus (Reduviidae: Emesinae). Proc.
Entomol. Soc. Wash. 77: 456-459.
Maldonado Capriles, J. 1976. The genus Heza
(Hemiptera: Reduviidae). J. Agric. Univ. Puerto
Rico 60: 403-433.
Maldonado Capriles, J. 1976. Two new genera
and some records of Miridae (Hemiptera) from
Panama. Proc. Entomol. Soc. Wash. 78: 6-11.
Maldonado Capriles, J. 1976. A remarkable Sten-
opoda from Jamaica (Hemiptera: Reduviidae).
Proc. Entomol. Soc. Wash. 78: 357-360.
Maldonado Capriles, J. 1976. Three new species
of Castolus and a key to the species (Hemiptera:
Reduviidae). Proc. Entomol. Soc. Wash. 78: 435—
446.
Maldonado Capriles, J. and J. Mir6 Mercado.
1977. The wing louse, Lipeurus caponis (L.)
(Mallophaga: Philopteridae) attacking poultry in
Puerto Rico. J. Agric. Univ. Puerto Rico 61:
309-310.
Maldonado Capriles, J. and A. Berrios. 1977.
The immature stages of Copestylum vacuum (F)
(Diptera: Syrphidae), a new record for Puerto
Rico. J. Agric. Univ. Puerto Rico 56: 395-398.
Maldonado Capriles, J. and M. Miro Mercado.
1977. The mite Ornithonyssus sylviarum (C. and
F) (Arachnida: Acarina: Macronyssidae) attack-
ing fowl in Puerto Rico. J. Agric. Univ. Puerto
Rico 56: 400—401.
Maldonado Capriles, J. and S. Medina Gaud.
1977. The ticks in Puerto Rico (Arachnida: Ac-
arina). J. Agric. Univ. Puerto Rico 61: 402—404.
Maldonado Capriles, J. 1977. Studies on Idiocer-
inae leafhoppers, XV. Busonia Distant and an al-
74.
76.
Uke
78.
-
80.
81.
82.
83.
84.
85.
86.
lied new genus from the Oriental Region (Cica-
dellidae: Idiocerinae). Pacific Insects 17: 491-—
501.
Maldonado Capriles, J. 1977. Studies on Idiocer-
inae leafhoppers, XVIII: Four new genera and
species from Guyana, South America (Homop-
tera: Cicadellidae). Proc. Entomol. Soc. Wash.
79: 358-366.
. Maldonado Capriles, J. 1977. Studies on Idiocer-
inae leafhoppers, XIX: A new genus from Suri-
nam, northern South America. Proc. Entomol.
Soc. Wash. 79: 605-608.
Maldonado Capriles, J. and T. H. Farr. 1977. As-
sassin bugs of Jamaica (Hemiptera: Reduviidae).
Slonea (Occas. Pap. Nat. Hist., Div. Inst. Jamai-
ca, Kingston) No. 2, 18 pp.
Maldonado Capriles, J. and A. Ortiz. 1978. The
allotype of Arenasella maldonadoi and change of
depository for two of Fennah’s holotypes (Ho-
moptera: Tropiduchidae and Issidae). Proc. En-
tomol. Soc. Wash. 80: 380-382.
Maldonado Capriles, J. 1980. The genus Jobertus
Distant, 1884 (Hemiptera: Miridae: Orthotyli-
nae). J. Agric. Univ. Puerto Rico 64: 304—309.
Maldonado Capriles, J. 1981. Una nueva especie
de Crassicornus Carvalho y clave para las es-
pecies del género (Hemiptera: Miridae: Phyli-
nae). Bol. Entomol. Venez. 1: 115-118.
Maldonado Capriles, J. 1981. A new Ghilianella
and a new saicine genus, Buninotus (Hemiptera:
Reduviidae) from Panama. J. Agric. Uniy. Puerto
Rico 65: 401—407.
Maldonado Capriles, J. 1981. Hemiptera collect-
ed by the Biospeological Expedition to Cuba by
the Academies of Cuba and Romania. Resultats
Expeditions Biospeologiques Cubano-Roumai-
nes a Cuba 3: 113-121.
Maldonado Capriles, J. 1981. A new Ghinallelia
from Cuba (Hemiptera: Reduviidae-Emesinae).
Resultats Expeditions Biospeologiques Cubano-
Roumaines a Cuba 3: 123-124.
Maldonado Capriles, J. and J. C. M. Carvalho.
1981. A new species of Platyscytus Reuter and a
key to the species of the genus (Hemiptera, Mir-
idae). Rev. Brasil. Biol. 41: 385-388.
Maldonado Capriles, J. and J. C. M. Carvalho
1981. New species of Guianella Carvalho from
Panama and a key to the species (Hemiptera, Mi-
ridae). Rev. Brasil. Biol. 41: 389-394.
Maldonado Capriles, J. 1982. Designation of the
senior types of Paraproba pendula Van Duzee,
1914, and a key to the North American species
of Paraproba (Hemiptera: Miridae). J. Agric.
Univ. Puerto Rico 66: 282-285.
Maldonado Capriles, J. and E. Abreu. 1982.
Speculations about phoresy by a deuteronymph
(Acarina) on Atractocerus brasiliensis (Coleop-
VOLUME 99, NUMBER 4
87.
88.
89.
90.
91.
92.
OB:
94.
3).
96.
Dl.
98.
PH).
100.
101.
tera: Lymexylonidae). J. Agric. Univ. Puerto
Rico 67: 514-517.
Henry, T. J. and J. Maldonado Capriles. 1982.
The four “‘ocelli’’ of the isometopine genus /so-
metocoris Carvalho and Sailer (Hemiptera: Mir-
idae). Proc. Entomol. Soc. Wash. 84: 245-249.
Carvalho, J. C. M. and J. Maldonado Capriles.
1982. Mirideos Neotropicais. CCXXXII: Descri-
¢ao de um novo genero e duas especies novas
(Hemiptera). Rev. Brasil. Biol. 42: 55-61.
Carvalho, J. C. M. and J. Maldonado Capriles.
1982. Neotropical Miridae, CCXXVIII: Genus
Lundiella Carvalho with descriptions of new spe-
cies (Hemiptera). Rev. Brasil. Biol. 42: 55-61.
Maldonado Capriles, J. and H. Brailovsky. 1983.
Mexican Reduviidae II: The genus Heza Amyot
and Serville, 1843 (Hemiptera: Harpactorinae).
Proc. Entomol. Soc. Wash. 85: 222-225.
Giacchi, J. C. and J. Maldonado Capriles. 1983.
A new species of Oncocephalus Klug, 1830 (Re-
duviidae: Stenopodainae) from Nicaragua. Carib.
J. Sci. 19: 61-63.
Maldonado Capriles, J. 1983. Five new species
of Eccritotarsus (Hemiptera: Miridae) from Pan-
ama and locality records for other species in the
genus. Carib. J. Sci. 19: 65-70.
Maldonado Capriles, J. and H. Brailovsky. 1983.
Estudio sobre los reduviidos méxicanos, Parte I:
Emesinae (Hemiptera: Reduviidae). Carib. J. Sci.
19: 71-90.
Maldonado Capriles, J. 1984. The genus Knigh-
tonia Carvalho and Drake (Miridae: Hemiptera:
Hyaliodini). J. Agric. Univ. Puerto Rico 68: 307—
Sill:
Maldonado Capriles, J. 1984. Studies on Idiocer-
inae leafhoppers, XVI: Tomopennis, a new genus
from Guyana and a key to the neotropical genera
of the subfamily. Carib. J. Sci. 20: 97-100.
Maldonado Capriles, J. 1983. Concerning new
and old species of Heza (Hemiptera: Reduvi-
idae). J. Agric. Univ. Puerto Rico 67: 407-418.
Maldonado Capriles, J. 1984. A new species of
Eurychilella (Hemiptera: Miridae). J. Agric.
Univ. Puerto Rico 68: 75-78.
Maldonado Capriles, J. 1984. The genus Afae-
nius (Coleoptera: Scarabaeidae) in Puerto Rico.
J. Agric. Univ. Puerto Rico 68: 111-112.
Maldonado Capriles, J. 1985. Studies on idiocer-
ine leafhoppers XX. Gressittocerus and Doli-
chopscerus, new genera from New Guinea (Ho-
moptera: Cicadellidae). Int. J. Entomol. 27: 270—
276.
Maldonado Capriles, J. 1985. Studies on idiocer-
ine leafhoppers XXI. Color variations of Jdios-
copus clypealis (Homoptera: Cicadellidae). Int. J.
Entomol. 27: 277-279.
Maldonado Capriles, J. and H. Brailovsky. 1985.
Mexican Reduviidae, III]. The genus Atrachelus
102.
104.
105.
106.
107.
108.
109.
110.
Lett
114.
IS).
116.
769
(Harpactorinae). J. Agric. Univ. Puerto Rico 69:
185-189.
Maldonado Capriles, J. 1986. Designation of lec-
totypes from New World reduviid syntypes in the
“Institut royal des Sciences naturelles de Bel-
gique”’ (Insecta: Hemiptera). Bull. Inst. R. Sci.
Belg. Entomol. 56: 41—44.
3. Maldonado Capriles, J. 1986. Concerning Cuban
Miridae (Insecta: Hemiptera). Carib. J. Sci. 22:
125-136.
Maldonado Capriles, J. 1986. The Reduviidae
(Hemiptera) of Puerto Rico: an annotated list. J.
Agric. Univ. Puerto Rico 70: 1-7.
Maldonado Capriles, J. 1986. The genus Pniron-
tis, the subinermis group of species (Hemiptera:
Reduviidae: Stenopodainae). J. Agric. Univ.
Puerto Rico 70: 9-18.
Maldonado Capriles, J. 1986. Two new species
of Apronius Stal with notes on the genus (Het-
eroptera: Reduviidae: Stenopodainae). J. New
York Entomol. Soc. 94: 174-179.
Maldonado Capriles, J. and S. Medina Gaud.
1987. Insectos daninos y beneficiosos de Puerto
Rico. (Manual de Laboratorio). Libreria Univer-
sal. Mayagiiez, PR. 200 pp.
Maldonado Capriles, J. 1987. Orbella, a new har-
pactorine genus (Hemiptera: Reduviidae). J.
Agric. Univ. Puerto Rico 71: 193-198.
Maldonado Capriles, J. 1987. New names in Ve-
ledella and Sastrapada. First description of male
of Lamotteus ornatus Villiers (Reduviidae). J.
Agric. Univ. Puerto Rico 71: 199-202.
Maldonado Capriles, J. 1987. Homalocoris punc-
tatus n. sp. and key to the species in the genus
(Reduviidae, Microtominae). J. Agric. Univ.
Puerto Rico 71: 249-253.
Abreu, E., S. Medina Gaud, and J. Maldonado
Capriles. 1987. New records of mites for Puerto
Rico. J. Agric. Univ. Puerto Rico 71: 399-402.
. Maldonado Capriles J. 1987. Lopodytes bequaer-
ti Schouteden and two allied new species (Insec-
ta: Reduviidae). Rev. Zool. Afr. 100: 395-398
(1986).
3. Maldonado Capriles J. 1988. A new species of
Systelloderes Blanchard from South Africa (He-
miptera: Enicocephalidae). Ann. Transvaal Mus.
34: 621-623.
Maldonado Capriles, J. 1988. New species and
redescriptions of South African Emesinae (He-
miptera: Reduviidae). Ann. Transvaal Mus. 34:
625-634.
Maldonado Capriles, J. 1988. A new species of
Chondrolophus and a key to the species (Insecta:
Hemiptera: Reduviidae). Bull. Ann. Soc. R. Belg.
Entomol. 124: 229-232.
Coscar6én, M. C. and J. Maldonado Capriles.
1988. Contribution to the knowledge of Rasahus
Amyot and Serville, 1843 (Hemiptera: Reduvi-
770
NA7/e
118.
file:
120.
12
122.
25:
126.
128.
1293
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
idae); description of a new species. J. Agric.
Univ. Puerto Rico 72: 247-250.
Carpintero, D. J. and J. Maldonado Capriles.
1988. Contributions to the knowledge of Amer-
ican Ectrichodiinae I. Notes about Rhiginia (He-
miptera: Reduviidae). J. Agric. Univ. Puerto Rico
72: 251-254.
Santiago Blay, J. A. and J. Maldonado Capriles.
1988. Observations on the true bugs Emesa te-
nerrima, a possible spider mimic, and Ghilianel-
la borincana (Hemiptera: Reduviidae: Emesinae)
from Puerto Rico. Proc. Entomol. Soc. Wash. 90:
369-372.
Santiago Blay, J. A. and J. Maldonado Capriles.
1989. A new structure on the hind legs of male
Monalocoris carioca Carvalho and Gomez (Het-
eroptera: Miridae). J. New York Entomol. Soc.
97: 479-482.
Carpintero, D. J. and J. Maldonado Capriles.
1990. Contributions to the knowledge of Amer-
ican Ectrichodiinae. II. Notes about Rhiginia and
Pothea (Hemiptera: Reduviidae). J. Agric. Univ.
Puerto Rico 74: 449—456.
Maldonado Capriles, J. 1990. Systematic Cata-
logue of the Reduviidae of the World. Special
Publ. Carib. J. Sci. 704 pp.
Maldonado Capriles, J. 1991. Additions and cor-
rections to “The Miridae of Puerto Rico” (In-
secta: Heteroptera). Carib. J. Sci. 27: 117-123.
3. Carpintero, D. J. and J. Maldonado Capriles.
1991. Contributions to the knowledge of Amer-
ican Ectrichodiinae. III]. The genus Cricetopareis
Breddin, 1903 (Hemiptera: Reduviidae). J. Agric.
Univ. Puerto Rico 75: 81-86.
. Maldonado Capriles, J. and P. W. Lozada Robles.
1991. A new species of Apronius Stal (Reduvi-
idae: Stenopodainae) from Peru. J. Agric. Univ.
Puerto Rico 75: 407—409.
Kun-Wei, H. and J. Maldonado Capriles. 1991.
Idiocerinae of Taiwan (Homoptera: Cicadelli-
dae). J. Taiwan Mus. 45: 1-14.
Maldonado Capriles, J. and J. A. Santiago-Blay.
1991. Classification of Homalocoris (Heterop-
tera: Reduviidae: Hammacerinae), with the de-
scription of a new species. Proc. Entomol. Soc.
Wash. 93: 703-708.
. Maldonado Capriles, J., E. Abreu, C. Negron,
and D. Rodriguez. 1991. Presencia del acaro Ty-
rophagus putrescientae (Schrank) (Acarina:
Acaridae) sobre un cadaver embalsamado. Sci-
ence-Ciencia (Ponce, PR) 17: 62—63.
Maldonado Capriles, J. and H. Brailovsky. 1992.
Mexican Reduviidae, IV: a new species of Cas-
tolus (Heteroptera: Harpactorinae). Carib. J. Sci.
28: 41-43.
Maldonado Capriles, J. 1992. A new genus and
two species from South Africa and a generic key
to the world apterous or micropterous Leistar-
130.
hile
i 3X6):
Sve
138.
139.
140.
141.
142.
chini (Heteroptera: Reduviidae: Emesinae). J.
Afr. Zool. 106: 527-533.
Maldonado Capriles, J. 1992. On a small collec-
tion of Reduviidae (Insecta: Heteroptera) from
Brazil. J. Agric. Univ. Puerto Rico 76: 9-14.
Froeschner, R. C. and J. Maldonado Capriles.
1992. A synopsis of bugs of Puerto Rico with
description of new species Melanaethus wolcotti
(Heteroptera: Cydnidae). J. Agric. Univ. Puerto
Rico 76: 177-185.
. Maldonado Capriles J. 1992. New host for Pseu-
dopteromalus calandrae (Howard) (Hymenop-
tera: Pteromalidae) in Puerto Rico. J. Agric.
Univ. Puerto Rico 76: 209.
3. Maldonado Capriles, J. and P. W. Lozada Robles.
1992. Key to the group of neotropical wasp-mi-
metic harpactorine genera and the description of
a new species (Hemiptera: Reduviidae). Proc.
Entomol. Soc. Wash. 94: 162-165.
. Maldonado Capriles, J. and J. A. Santiago-Blay.
1992. A new species of the neotropical genus
Sphaeridops Amyot & Serville, 1843 (Sphaeri-
dopinae: Reduviidae). Proc. Entomol. Soc. Wash.
94: 508-511.
5. Maldonado Capriles, J. 1993. A new harpacto-
rine genus from French Guiana, northern South
America. (Heteroptera: Reduviidae). Carib. J.
Sci. 28: 38—40.
Maldonado Capriles, J., J. A. Santiago-Blay, and
G. O. Poinar, Jr. 1993. Apicrenus fossilis gen. &
sp. n. (Heteroptera: Reduviidae: Apiomerinae)
from Dominican amber (lower Oligocene—upper
Eocene). Entomol. Scand. 24: 139-142.
Maldonado Capriles, J. 1993. Four new enico-
cephalids from Africa (Heteroptera: Enicoce-
phalidae). J. Afr. Zool. 107: 57-63.
Maldonado Capriles, J., J. A. Santiago-Blay, and
G. O. Poinar, Jr. 1993. Paleoploiariola venosa, a
new fossil Emesinae (Heteroptera: Reduviidae)
genus and species from Dominican amber. J.
Agric. Univ. Puerto Rico 77: 95-100.
Maldonado Capriles, J. 1993. A newly discov-
ered homonym in Ectomocoris (Heteroptera: Re-
duviidae: Peiratinae). Proc. Entomol. Soc. Wash.
95: 128.
Maldonado Capriles, J. and D. J. Carpintero.
1993. Redescription of the harpactorine genus
Sosius Champion 1899, with the description of a
new species (Heteroptera: Reduviidae). Proc. En-
tomol. Soc. Wash. 95: 223-227.
Maldonado Capriles, J. 1993. A new species of
Ghilianella and the female of Ghinallelia clavi-
ventris (Bergroth) (Heteroptera: Reduviidae:
Emesinae). Proc. Entomol. Soc. Wash. 95: 558—
561.
Maldonado Capriles, J. 1993. Redescription of
Nesocastolus nigricornis (Fracker and Bruner)
and notes on the genus (Heteroptera: Reduviidae:
VOLUME 99, NUMBER 4
143.
144.
145.
146.
147.
148.
149.
150.
LESIIE
152:
Harpactorinae). Proc. Entomol. Soc. Wash. 95:
562-563.
Maldonado Capriles, J. 1994. Pnirontis grandis
n. sp. (Heteroptera: Reduviidae: Stenopodainae).
Proc. Entomol. Soc. Wash. 96: 713-716.
Maldonado Capriles, J. 1994. A new species of
Seridentus Osborn, 1904 and a key to the species
(Heteroptera: Reduviidae: Stenopodainae). Proc.
Entomol. Soc. Wash. 96: 717-722.
Maldonado Capriles, J. 1994. Hemiptera-Heter-
optera. pp. 307-311. Jn: C. Juberthie et V. Decu
(eds.). Encyclopaedia Biospeologica. Societe de
Bioespeologie. Bucarest, Roumanie. Tome I, pp.
1-xiit+7—311.
Maldonado Capriles, J. 1995. A new Ctenotra-
chelus and notes about other species in the genus
(Heteroptera: Reduviidae: Stenopodainae). Carib.
J. Sci. 31: 141-147.
Maldonado Capriles, J. and G. O. Poinar, Jr.
1995. Stomatomiris, a new fossil mirid genus in
Dominican amber (Heteroptera: Miridae). Carib.
J. Sci. 31: 281-283.
Maldonado Capriles, J. 1995. Notes about the
Old World genus Hexamerocerus Reuter (Het-
eroptera: Reduviidae: Ectrichodiinae). Proc. En-
tomol. Soc. Wash. 97: 112-116.
Maldonado Capriles, J. and G. O. Poinar, Jr.
1995. Reuteroscopus carvalhoi n. sp., a new fos-
sil plant bug (Heteroptera: Miridae: Phylinae).
Proc. Entomol. Soc. Wash. 97: 346-349.
Maldonado Capriles, J. 1995. Notes about Ves-
cinae, a key to the world genera, and description
of two new Chopardita (Heteroptera: Reduvi-
idae). Proc. Entomol. Soc. Wash. 97: 626-633.
Maldonado Capriles, J. 1995. New nearctic spe-
cies of Oncocephalus Klug (Heteroptera: Redu-
viidae: Stenopodainae). Proc. Entomol. Soc.
Wash. 97: 791-798.
Carpintero, D. J. and J. Maldonado Capriles.
Vs.
1996. Diagnostic characters and key to the gen-
era of American Ectrichodiinae (Heteroptera, Re-
duviidae). Carib. J. Sci. 32: 125-141.
3. Maldonado Capriles, J. 1996. Reduviidae (Het-
eroptera) collected by fogging the forest canopy
in Peru. Proc. Entomol. Soc. Wash. 98: 233-236.
Maldonado Capriles, J. 1996. New taxa and key
to the tribes and genera in Tribelocephalinae Stal
1866 (Heteroptera: Reduviidae). Proc. Entomol.
Soc. Wash. 98: 138-144.
5. Maldonado Capriles, J. and P. H. van Doesburg.
1996. A new genus and species of Emesinae
(Heteroptera: Reduviidae) from Suriname. Zool.
Meded. 70: 93-97.
Maldonado Capriles, J., J. A. Santiago-Blay, and
G. O. Poinar, Jr. 1996. Paralienates hyalinus n.
gen., n. sp. (Enicocephalidae: Heteroptera): A
fossil gnat bug from Dominican amber (lower
Miocene—upper Eocene). J. Agric. Univ. Puerto
Rico 80: 47-54.
Maldonado Capriles, J. 1996. The status of insect
alpha taxonomy in Puerto Rico after the Scien-
tific Survey. pp. 201-216. In: J. C. Figueroa Co-
l6n (ed.). The Scientific Survey of Puerto Rico
and the Virgin Islands. An Eighty-Year Reas-
sessment of the Islands’ Natural History. Vol. 76.
273 pp.
Jorge A. Santiago-Blay, Department of
Ecology and Evolution, 1101 East 57th
Street, The University of Chicago, Chicago,
IL 60637, U.S.A.; Thomas J. Henry, Sys-
tematic Entomology Laboratory, PSI, Ag-
ricultural Research Service, U.S. Depart-
ment of Agriculture, c/o National Museum
of Natural History, MRC-168, Washington,
DE 205607 USA’ Nand A"Gs Wheeler Ir;
Department of Entomology, Clemson Uni-
versity, Clemson, SC 29634, U.S.A.
154.
156.
STE
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 772-777
SOCIETY MEETINGS
1016th Regular Meeting—January 9, 1997
The 1016th Regular Meeting of the En-
tomological Society of Washington (ESW)
was called to order at 8:02 pm by President
M. Alma Solis on January 9, 1997, in the
Waldo Schmitt Room of the National Mu-
seum of Natural History, Washington, DC.
There were 9 members and 8 guests pres-
ent. The minutes of the December 5, 1996,
meeting were read by Acting Recording
Secretary, Dawn Southard. This included a
correction to a correction to the November
1996 minutes. It was stated that one of the
visitors to the December ’96 ESW meeting,
Manual Balcazar, had already earned his
Ph.D., and was at the Instituto de Biologia,
UNAM, Mexico; not still a student at the
University of Florida. The minutes were
then approved as corrected.
There were no officers’ reports, no com-
mittee reports, and no old business. Presi-
dent Solis asked if anyone had notes or ex-
hibits. Warren Steiner showed and passed
around a new book from South Africa,
which had been given to him by our eve-
ning’s speaker, Dr. Bellamy, entitled: Revi-
sion of the Tribe Caenocrypticini, Coleop-
tera: Tenebrionidae: Tentyriinae by S. En-
drody-Younga, Transvaal Museum Mono-
graph No. 11. Oct. 1996, ISBN 0-907990-
177. David Furth showed and passed around
a new book, entitled: An Inordinate Fond-
ness for Beetles by Arthur V. Evans and
Charles L. Bellamy [our evening’s speaker],
1996, published by Henry Holt & Co.; ISBN
0-8050-3751-9. David Furth also showed a
new T-shirt covered with illustrations of in-
sects (mainly beetles) of the world. M. Alma
Solis described a children’s book she had re-
cently read called “‘Miss Spider’s Tea Party.”
David Furth, ESW Program Chair, intro-
duced our speaker for the evening: Dr.
Charles L. Bellamy, Coleoptera Depart-
ment; Transvaal Museum; PO. Box 413;
Pretoria 0001, South Africa. Dr. Bellamy
earned his B.S. and M.S. from California
State University-Long Beach, under Dr.
Sleeper; and his D.Sc. from the University
of Pretoria, South Africa. His presentation,
entitled ‘““‘Beetle Collecting in Southern Af-
rica,” gave a brief overview of the habits,
physiography, and peoples of southern Af-
rica (especially Malawi, South Africa, and
Zimbabwe), followed by a general discus-
sion of beetle diversity. Many excellent
slides were shown of the region’s diverse
habitats, geological features, vegetation,
megafauna, and a number of more com-
monly encountered beetle species.
Five visitors were introduced: Dennis
Hollier, manager of a Georgetown area ho-
tel and husband of our acting recording sec-
retary, Dawn Southard; Steve Lingafelter, a
new staff member of the Systematic Ento-
mology Laboratory; Dr. Alfonso Alonso, a
former student of Dr. Brower at the Uni-
versity of Florida; Dr. Leeanne E. Alonso,
who earned her Ph.D. from Harvard fol-
lowed by a postdoctoral fellowship study-
ing ant-plant interactions at the University
of Oklahoma; and Dr. John W. Brown, who
earned his Ph.D. from the University of
California, Berkeley, and is a new System-
atic Entomology Laboratory staff member.
President Solis adjourned the meeting at
9:10 pm. Refreshments were provided by
Jil Swearingen and Warren Steiner.
Respectfully submitted,
Harold J. Harlan,
Recording Secretary
1017th Regular Meeting—
February 6, 1997
The 1017th Regular Meeting of the En-
tomological Society of Washington (ESW)
was called to order at 8:01 pm by President
M. Alma Solis on February 6, 1997, in the
Waldo Schmitt Room of the National Mu-
seum of Natural History, Washington, DC.
There were 14 members and 4 guests pres-
VOLUME 99, NUMBER 4
ent. The minutes of the January 9, 1997,
meeting were read by Recording Secretary,
Harold Harlan. David Furth pointed out that
Dr. John W. Brown, a visitor at the January
9, 1997 meeting, was working at the USDA's
Systematic Entomology Laboratory, not at
the Smithsonian; and that he had earned his
Ph.D. at U.C. Berkeley, not at San Diego.
The spelling of one visitor’s name was cor-
rected to ‘“‘Leeanne’’ Alonso. The minutes
were approved after those corrections.
There were no officers’ reports, no com-
mittee reports, and no old business. Presi-
dent Solis asked if there were any visitors
or new members. Dr. John W. Brown be-
came a new member since the January °97
meeting. President Solis asked if anyone
had notes or exhibits. David Smith showed
the new Memoir of the ESW (No. 18), Con-
tributions on Diptera, dedicated to Willis
W. Wirth and edited by Wayne Mathis and
William L. Grogan, Jr. He also briefly dis-
cussed the confusion which might arise be-
cause the date inside was 1996, but the
mailing date was January 1997. He next
showed and passed around a nice picture of
a large (25 mm), north temperate sawfly,
Cimbex americana Leach, from Washing-
ton State. Anyone sighting such specimens
around here please let him know. It is an
elm sawfly, also found on willow and
weeping willow. He further commented that
we should be alert for sawfly activity soon.
His earliest record of sawflies in this area
has been February 17. President Solis re-
ferred us all to Georgia Brown’s restaurant
on 16th St. (just south of K St.) in NW DC,
which has sawflies (about 3-in. long) paint-
ed in gold on its front doors and windows.
David Furth showed and passed around
three new books: Spineless Wonders, by
Richard Coniff (our scheduled April 1997
speaker), 1996, published by Henry Holt &
Co, NY, ISBN: 0-8050-4218-0; and two
volumes by Gordon M. Nishida & Joann
M. Tenorio—What’s Bugging Me?, 1995,
Univ. of HI Press, ISBN: 0-8248-1742-7;
and What Bit Me?, 1993, Univ. of HI Press,
ISBN: 0-8248-1492-4. Ed Saugstad showed
Ve
and passed around a copy of a pamphlet
from Japan entitled: ‘‘Techno Insect,”’ [Vol.
26 (No. 6): 8-34, June 1993], which high-
lighted the known and potential value of in-
sects from a technologic point of view.
David Furth announced that next month’s
speaker will be Eric Grissell, of SEL,
USDA. He then introduced this evening’s
speaker, our Past President, Dr. Ralph Eck-
erlin: Northern Virginia Community Col-
lege, Annandale, VA. Ralph earned his B.S.
in Biology from Rutgers, and his M.S. in
Zoology from the Univ. of Miami, FL, with
research emphasis on flukes. He worked 7
years for American Cyanamid, then
‘“‘wormed”’ his way back into pursuing a
Ph.D., at the Univ. of CT with research on
nematodes. He shared an office with Harry
Painter for several years, and they collab-
orated on most of the research included in
this evening’s talk. His presentation, enti-
tled: ‘“‘Ectoparasite Potpourri,’ summarized
several years of research on ectoparasites.
Much of it related to fleas, their biology and
zoogeography. Large changes in relative
proportions of nest-inhabiting fleas of
southern flying squirrels were discussed.
These changes were associated with time of
year, temperature, and elevation. Orchopeas
howardii (Baker) was dominant at lower el-
evations and summer; but was replaced by
Opisodasys pseudarctomys (Baker) at high-
er elevations (=2000 ft.) in winter. Ob-
served distributions of some of these flea
species suggest ““mountaintop speciation”
along the Appalachians. Later studies on
ticks and on leptinid beetles (Coleoptera:
Leptinidae) were also discussed. Some very
good slides were included of recent ecto-
parasite studies Dr. Eckerlin had done in
Central America with local national and
U.S. collaborators.
The meeting was adjourned at 9:07 pm.
Refreshments were provided by M. Alma
Solis.
Respectfully submitted,
Harold J. Harlan,
Recording Secretary
774
1018th Regular Meeting—March 6, 1997
The 1018th Regular Meeting of the En-
tomological Society of Washington (ESW)
was called to order at 8:04 pm by President
M. Alma Solis on March 6, 1997, in the
Waldo Schmitt Room of the National Mu-
seum of Natural History, Washington, DC.
There were 18 members and 9 guests pres-
ent. The minutes of the February 6, 1997,
meeting were read by Recording Secretary,
Harold Harlan.
There were no officers’ reports and no
old business. Holly Williams reported, on
behalf of the Membership Chair, there are
seven prospective new members this month
(March 1997). They are: Carol DiSalvo;
Warren G. Abrahamson; Marion Kotrba;
Alexander S. Konstantinov; Steve Linga-
felter; Steve W. Chordas, II]; and Diane M.
Calabrese. David Furth reported, on behalf
of the Executive Committee, that regular
ESW meetings will begin earlier: April’s
meeting will begin at 7:30 pm, and subse-
quent meetings may be moved up to start
at 7:00 pm. No new members were present.
David Furth introduced a visitor, Andy
Brower, a Post-doctoral Fellow studying
Lepidoptera at the USNM. President Solis
then asked for presentation of any notes or
exhibits. Dr. Louis J. Pribyl, a research col-
laborator in the Department of Paleobiolo-
gy, Smithsonian, showed and briefly dis-
cussed fossil insects he had displayed at the
back of the room. These were Eocene
(Green River) fossil insects from Piceance
Creek Basin, CO. He invited any experts to
assist in identifying these specimens and
provided an information sheet and invita-
tion to interested parties (copies are on a
rear table). Please review one of these and
take a copy with you for further review or
follow-up. David Furth showed and placed
on display three new publications of pos-
sible interest to members: Insects of the
Great Lakes Region, by Gary Dunn, 1996;
published by the Univ. of Michigan Press,
Ann Arbor; ISBN (hard cover): 0-472-
09515-3; ISBN (paperback): 0-472-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
06515-7. Nomina Insecta Nearctica: A
Checklist of the Insects of North America;
Diptera, Lepidoptera, and Siphonaptera.
Vol. 3, 1996; compiled by R. W. Poole &
R. E. Lewis; by Entomological Information
Services, Rockville, MD; ISBN (series):
1-889002-00-3; ISBN [Vol. 3]: 1-889002-
03-8; and a pamphlet-style report on the
Permits Workshop, San Diego, CA, 29-31
January 1997. Gabriela Chavarria showed
and passed around a book entitled: The
Conservation of Bees, by A. Matheson, S.
L. Bachmann, C. O’ Toole, P. Westrich, & I.
H. Williams, 1996; by Academic Press, San
Diego, CA; ISBN: 0-12-479740-7. Ralph
Eckerlin showed and placed on display a
small metal can of DDT powder, which he
had found among the possessions of the late
Dr. Robert Traub, a long-time ESW mem-
ber and a well-known flea specialist. David
Furth next read a limerick related to the
speaker’s subject matter; and then intro-
duced the evening’s speaker, Dr. Eric Gris-
sell, SEL, USDA. Dr. Grissell earned his
B.S., M.S., and his Ph.D. from U.C. Davis.
He received his Ph.D. in 1972, studying the
origins of western species of Torymus in the
USS.
Biological diversity is very topical, and
within the biological community it is great-
ly pondered, debated, and written about. Bi-
ologists generally agree that a biodiversity
“‘crisis’’ exists. Each biologist’s reaction to
the dilemma may be focused on the per-
spective from his or her own specialized
point of view. Biologists are not the only
students of biodiversity, and we may fail to
heed the notion that dissidents to the con-
cept of biodiversity exist and that they
voice opinions which most mainstream bi-
ologists find appalling. To the overwhelm-
ing majority of “the public,’ the concept
of biodiversity is at best an acknowledg-
ment that life exists and at worst a violent
aversion to any lifeform that enters their
personal space.
Dr. Grissell illustrated some notions
about biodiversity that might seem alien to
most of us, and briefly explored the notion
VOLUME 99, NUMBER 4
that biodiversity is not a universally ac-
cepted paradigm within the scientific com-
munity (or in some cases what passes as the
scientific community). He presented a min-
imum number of factoids concerning Hy-
menoptera and discussed aspects of their
biodiversity that may be foreign to most of
us. And he presented some notions of pub-
lic perception concerning Hymenoptera and
insects and suggested that it is up to biolo-
gists to help make biodiversity a household
word of some importance to the majority of
the public. This is still quite a challenge.
The speaker’s approach reflected his per-
spective as a systematist with a lifelong in-
terest in the environment and its biodiver-
sity. Because he is a specialist in the Hy-
menoptera, he used examples from that
group to illustrate his points. Although that
may have appeared to be an insular choice,
he was not entirely convinced that other
groups of organisms, for example the bee-
tles, could provide more suitable examples
or make more sense.
The meeting was adjourned at 9:09 pm.
Refreshments were provided by Harold
Harlan.
Respectfully submitted,
Harold J. Harlan,
Recording Secretary
1019th Regular Meeting—April 3, 1997
The 1019th Regular Meeting of the En-
tomological Society of Washington (ESW)
was called to order at 7:30 pm by President
M. Alma Solis on April 3, 1997, in the Wal-
do Schmitt Room of the National Museum
of Natural History, Washington, DC. There
were 13 members and 15 guests present.
The minutes of the March 6, 1997, meeting
were read by Recording Secretary, Harold
Harlan.
There were no officers’ reports and no
old business. Holly Williams reported, on
behalf of the Membership Chair, that three
new members are Kipling Will, Marty Con-
don, and Lynn E. Carroll.
Under new business, President Solis an-
TS
nounced that the Executive Committee had
met and decided how to handle the excess
archived publications of the ESW. David
Furth has volunteered to contact some book
dealers as a possible partial solution to this
problem. President Solis asked for presen-
tation of any notes or exhibits. David Furth
showed and placed on display two publi-
cations of possible interest to members:
Thermal Warriors; Strategies of Insect Sur-
vival by Bernd Heinrich, 1996; published
by Harvard Univ. Press, Cambridge, MA.;
ISBN (hard cover): O0-674-88340-3; and
The Hot-Blooded Insects: Strategies and
Mechanisms of Thermoregulation; by
Bernd Heinrich, 1993; published by Har-
vard Univ. Press, Cambridge, MA; ISBN
(hard over): 0-674-40838-1.
David Furth announced that the Ento-
mology Dept., Smithsonian Institution, has
a lot of excess sheets of old-fashioned (=
“‘real’’) papered cork, each about 10” X 20.”
He showed one such sheet. Interested per-
sons should contact him. He also described
a computer mouse pad which is designed to
look like a ladybird beetle. Warren Steiner
announced that the annual ESW Banquet
will be held Tuesday evening, June 3rd,
1997, at the Uniformed Services Univ. din-
ing room. The speaker will be Mark Moffet,
a well known photo-journalist, from the
Museum of Comparative Zoology, Harvard
University. David Furth next introduced our
speaker for the evening, Richard Conniff, a
widely known nature journalist.
Richard Conniff is from Deep River, CT.
He is a prolific and accomplished writer and
producer of nature (especially invertebrate
animals) topics. He became interested in
nature relatively late in life. He earned a
B.S. in English from Yale (Class of 1973)
and worked as a journalist for a daily news-
paper until he found ‘“‘leeches were more
interesting than politicians.’”’ His subse-
quent works usually detail natural subjects
and often compare them to our frequently
‘unnatural’? everyday human lives. He has
written for Time, Smithsonian, Atlantic
Monthly, The New York Times, and Nation-
776
al Geographic, among others. He has nar-
rated and written and produced television
shows for the ‘National Geographic Ex-
plorer,”’ the Discovery Channel, and other
network nature specials. His script for one
National Geographic show, “‘Ocean Drift-
ers’’ was nominated for an Emmy Award
in 1993. He has recently written a book,
Spineless Wonders, published by Henry
Holt & Co. (ISBN: 0-8050-4218-0), from
which he drew much of his talk. He showed
some excellent slides of various aspects of
four main invertebrate groups: fire ants, ta-
rantulas, fleas, and moths. Both his verbal
and pictoral illustrations were often strik-
ing. For example, he compared the repro-
ductive function of a fire ant queen to be
roughly equivalent to ‘‘a 120-lb. woman
giving birth to 500,000 eight-pound babies
a year’’; he described tarantulas as “‘spiders
in a mink coats”’ and stated that they are
the “‘dominant predators of the rain forest’;
he noted that men once wooed women with
“poetry about their fleas’’; and he described
and showed examples of how moths’ eggs
mimic bird droppings and their larvae mim-
ic twigs and lichens on trees. The talk was
fascinating and concurrently very entertain-
ing. A few copies of his book were avail-
able for purchase and he signed those pur-
chased after the meeting.
The meeting was adjourned at 8:49 pm.
Refreshments afterward were provided by
John Brown.
Respectfully submitted,
Harold Harlan,
Recording Secretary
1020th Regular Meeting—May 1, 1997
The 1020th Regular Meeting of the En-
tomological Society of Washington (ESW)
was called to order at 7:53 pm by President
M. Alma Solis on May 1, 1997, in the Wal-
do Schmitt Room of the National Museum
of Natural History, Washington, DC. There
were 16 members and 10 guests present.
The minutes of the April 3, 1997, meeting
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
were read by Recording Secretary, Harold
Harlan.
Under Officers’ reports, President Solis
announced that the former Membership
Chair, David Adamski, resigned that posi-
tion and John Brown has been appointed as
the new Membership Chair. Warren Steiner
announced that Mark Moffett, our planned
banquet speaker, has been given, with very
little notice, an important assignment in Viet-
nam and will not be available on June 3.
However, Bernd Heinrich, Zoology Depart-
ment, University of Connecticut, has agreed
to speak at our banquet. His tentative title
will be something like ““Adventures with
Hot and Cold Dung-Rolling Beetles.”
There were no other officers’ reports or
committee reports. John Brown, Member-
ship Chair, read names of two new mem-
bers: Patricia Jimenez Guarda and Imelda
Martinez Morales. Two visitors were intro-
duced. David Furth introduced Dr. Lee, a
visiting scientist from Andong University,
Republic of Korea, working on chrysome-
lids. Harry Painter introduced his wife, Au-
drey.
David Furth announced that Lee Goff, a
well-known Forensic Entomologist and
Acarologist from the University of Hawaii,
will be visiting this area and will present a
talk on Forensic Entomology, here in the
Waldo Schmitt Room, at 1:00 pm Tuesday,
May 20, 1997. Everyone is invited.
President Solis asked for presentation of
any notes or exhibits. Paul Spangler showed
and passed around a colored picture of two
lucanid beetles with mites on them (one
was almost completely covered). He also
showed and passed around two fossil bee-
tles: one was a scarab from Ecuador; the
other was probably a dyticid from Wash-
ington. David Furth showed and placed on
display four publications of possible inter-
est to members: 1. Ecology of Coccinellidae
by I. Hodek and A. Honek, 1996; published
by Kluwer Academic Publishers, Dor-
drecht, ISBN (hard cover): 0-7923-4177-5;
2. Canopy Arthropods, ed. N. E. Stork, J.
Adis, and R. K. Didham, 1997; published
VOLUME 99, NUMBER 4
by Chapman and Hall, London, ISBN (hard
cover): 0-412-74900-9; 3. Keys to the In-
sects of the European Part of the USSR.
(Translation), Vol. If], Hymenoptera, Part 5.
ed., G. S. Medvedev, 1995; Science Pub-
lishers, Inc. Published by the Institute of
Zoology, Academy of Sciences of the
USSR. No. 147, ISBN: 1-886106-24-X,
printed at Baba Barkha Nath Press, New
Delhi: and 4. The Carrion Beetles (Cole-
optera: Silphidae) of Nebraska by Brett C.
Rattcliffe, Bull. of the University of Ne-
braska State Museum, Vol. 13, 1996; pub-
lished by the University of Nebraska State
Museum, ISBN: 0093-6812.
David Furth next introduced our speaker
for the evening, Mila Coca, who is nearing
the end of a two-year post-doctoral fellow-
ship provided by her own Spanish govern-
ment. Mila had earned her Ph.D. in Madrid,
Spain, studying the phylogeny of scarabae-
id beetles. She will soon return to a position
in the National Museum of Spain. She has
been studying Phyllophaga spp. of the U.S.
under Terry Erwin at the Smithsonian In-
stitution.
Her talk addressed relationships of taxa
within the Melolanthini (‘“‘May”’ or “‘June”’
beetles) of Europe and the U.S. She com-
pared and contrasted their morphologies
and, to some extent, their behavior. Larvae
(subterranean forms) of certain groups are
very territorial, and often kill congeneric or
conspecific intruders. Mila discovered and
used many new morphological characters of
adult beetles’ genitalia, sterna, and even in-
ternal structures in her studies. She noted
that scanning electron micrographs can be
very helpful. Her results may help more ac-
curately identify serious agricultural pests
in Spain.
The meeting was adjourned at 8:28 pm.
Refreshments afterward were kindly pro-
vided by Paul and Phyllis Spangler.
Respectfully submitted,
Harold Harlan,
Recording Secretary
ela
1021st Regular Meeting—June 3, 1997
The Annual Banquet of the Entomolog-
ical Society of Washington (ESW) was held
Tuesday, June 3, 1997, in the dining room
of the Uniformed Services University of the
Health Sciences (USUHS), Bethesda, MD,
at 7:00 pm. The Maryland Entomological
Society (MES) shared in sponsoring this
event.
The evening began with a cash bar at 6:00
pm, accompanied by a lot of good conver-
sation and fellowship. The meal was served
at 7:00 pm followed by a brief program and
the presentation by our invited speaker.
President M. Alma Solis introduced David
Furth, Master of Ceremonies for the eve-
ning. Very brief remarks were given by:
Bob Platt, representing MES; Harold Har-
lan, representing USUHS; and Warren
Steiner, ESW President-Elect. David Furth
introduced our speaker, Dr. Bernd Heinrich,
Department of Zoology, University of Ver-
mont.
Dr. Heinrich’s talk, on ““Adventures with
Hot and Cold Dung-ball Rolling Beetles,”
included comparisons of different strategies
by dung-ball rolling beetles for finding and
exploiting various kinds and sizes of dung.
Those species with higher body tempera-
tures seemed to have an edge in finding and
beginning to process dung quickly. This
could be very critical where dung is scarce
or where it may dry out very quickly, thus
becoming unusable. His talk also addressed
different burying techniques and mating
habits. He showed many fine slides of the
beetles in action.
David Furth thanked our speaker for his
excellent presentation, thanked everyone
for attending (more than 70 people were
present) and wished all a safe trip home.
Alma Solis echoed those wishes, then
closed the banquet at about 9:45 pm.
Respectfully submitted,
Harold Harlan,
Recording Secretary
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, p. 778
RECENT PUBLICATIONS OF THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON
Memoir No. 17
Contributions on Hymenoptera and Associated Insects,
Dedicated to Karl V. Krombein. Edited by B. B. Norden and A. S. Menke.
216 pp. 1996 (cost: $25.00)
A Festschrift in recognition of Dr. Karl V. Krombein, Smithsonian Institution,
Washington, D.C. A collection of 22 invited papers, mostly on Hymenoptera, from Dr.
Krombein’s colleagues around the world. Biographic highlights and a bibliography of Dr.
Krombein’s publications by Beth B. Norden are presented.
Memoir No. 18
Contributions on Diptera, Dedicated to Willis W. Wirth.
Edited by Wayne N. Mathis and William L. Grogan, Jr.
297 pp. 1997 (cost: $25.00)
A memorial volume dedicated to D. Willis W. Wirth, eminent Dipterist and long-time
employee of the Systematic Entomology Laboratory, U.S. Department of Agriculture,
Washington, D.C. A collection of invited papers, mostly on Diptera, prepared by Dr.
Wirth’s colleagues around the world.
Memoir No. 19
Monograph of the Stilt Bugs, or Berytidae (Heteroptera), of the
Western Hemisphere. By Thomas J. Henry.
149 pp. 1997 (cost: $18.00)
Western Hemisphere Berytidae include 3 subfamilies, 13 genera, and 53 species. Keys
and numerous illustrations are provided for identification, and information on host plants,
feeding habits, and economic importance are provided. An important, comprehensive work
on this family.
Miscellaneous Publication
A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera)
Second Edition, Revised.
By E. Eric Grissell and Michael E. Schauff.
87 pp. 1997 (cost: $15.00)
This is a revised edition of the 1990 handbook on chalcids by the same authors. The
content is updated with the numerous advances that have appeared since 1990. Pictorial
keys and discussions of the characteristics, biology, distribution, and literature of each
family as well as collecting and mounting techniques and morphology make this a most
useful introduction and overview of the superfamily.
PROC. ENTOMOL. SOC. WASH.
99(4), 1997, pp. 779-784
PROCEEDINGS
of the
ENTOMOLOGICAL SOCIETY
of
WASHINGTON
Volume 99
OFFICERS FOR THE YEAR 1997
President M. Alma Solis
President-Elect Warren E. Steiner, Jr.
Recording Secretary Harold Harlan
Corresponding Secretary Hollis B. Williams
Treasurer Michael G. Pogue
Program Chair David G. Furth
Membership Chair David Adamski, John W. Brown
Custodian Gary L. Miller
Editor David R. Smith
Past President Ralph P. Eckerlin
Published by The Society
WASHINGTON, D.C.
1997
TABLE OF CONTENTS, VOLUME 99
ARTICLES
ABRAHAMSON, WARREN G.—See MELIKA, GEORGE ...................2. 0c cee cece ceeeeeee 560
ABRAHAMSON, WARREN G.—See MELIKA, GEORGE .................-.. 2-02 ccccecescceenes 666
ADAMSKI, DAVID and BERNARD LANDRY—Review of the Blastobasinae (Lepidoptera:
Gelechioidea: Coleophoridae) of the Galapagos Islands ................ 020.022 eeeeee cece eee eee 348
7N AION el DSS 2 0 Ba eS es on See Be AO nis eA ac RAB Spee Sa BAe Aga SOUS DGOCOOOr 143
BAL CIUNAS MOSERH K= SeeiGAGNEY RAYMOND IP ok ee ees 312
BARRE RAG ERINESTO—Scel BRAMBOMSRGYS EVAR YS we ticee loeceeielele ole etalcloel-miclaiale re eteielelalistereyeeieletese= 257
BATRA, SUZANNE W. T.—Bionomics of Lasioglossum (Evylaeus) matianense (Blithgen)
(Hymenoptera: Halicitidae), the predominant pollinating bee in orchards at high altitude in
the Great Himalaya of Garhwal, U.P., India ................. 2.0. .c cece cece cence eee tee eee e ees cenees 162
BELLOWS, TOM—See SWEARINGEN, MICHEL .................... ccc cece eee eeeeteee cess eee eees 513
BOHART, RICHARD M.—A review of the genus Hoplisoides Gribodo (Hymenoptera: Sphe-
cidaes Gonytimi)) im eNorth Amen Cate renen casei cscs ores stelote as a etetoinla el ete tele = -ieinte elelelole o!=ielaiela s/-loreeiale mim 645
780 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
BOYDSTON,, L.—See: PINE; KK.SS 4.0. tien soaietae cities see ee eee See Orne FER eee ee Ene 143
BRAILOVSKY, ARR Y—See SEATERS JAMES s Agere reenter e ee oee ee ene eer eee 37
BRAILOVSKY, HARRY and ERNESTO BARRERA—Redescription of the subgenus Hygia
(Eucolpura) Breddin (Hemiptera: Coreidae: Colpurini), with the description of two new spe-
cies; and akey to the known! Species #.-n ea ote eer ene ee eee ane eee eee eee nee e eee USSTI
BUENO-SORIA, JOAQUIN and SILVIA SANTIAGO-FRAGOSO—Studies in aquatic insects
XII: descriptions of nineteen new species of the genus Ochrotrichia Mosely (Trichoptera:
Hy droptilidac) irom \Viexicoands Central Aime Giclee eee ce eee eee eee 359
BURKES HORACE IR Scev ONES ROB ERG W saree eeeeee oeeeee eee eae eee eee eee eee 705
BURROWS, DAMIEN W.—Seée\GAGNE, RAYMOND J. 20.00. ..).c0:c. ccc decdevececveveceenese 312
BYERS, GEORGE W.—Four puzzling new species of Mecoptera ....................00eeeeee00ee 681
CONDON, MARTY, ANN TRUELOVE, and LYNN MATHEW—Sexual dimorphism in
mouthparts of Blepharoneura Loew (Diptera: Wephritidae)) s...--- 22-262 +2222 eee eee seeeeeeieee 676
COSCARON, MARIA DEL C. and JUAN J. MORRONE—Cladistics and biogeography of the
assassin bug genus Melanolestes Stal (Heteroptera: Reduviidae) ......................2...-.000- PP)
DEBARR?G.—See) TURGEON, Ji isc seeninguuce sects +s merase caer! anya sala aceon eee Eee Eee 608
DE VRUPIMARK—See ROBINSONT HAROLD pe eeee ee eee ee eee eee reece eae eee ee Eres 477
DUSSE, KATHLEEN and L. E. HURD—Food limitation reduces body length in mantid
nymphs, Tenodera sinensis Saussure (Mantodea: Mantidae): implications for fitness ......... 490
EASTON, EMMET R. and WING-WAH PUN—Observations on some Hemiptera/Heteroptera
of: Macau, ‘Southeast'Asiae sis. sneer ee cae erate eee ie ain eran ee ee 574
EVANS, HOWARD E. and STUART M. FULLERTON—Report on a collection of Bethylidae
(iiymenoptera) itiromicentrall hl onidageper en --pe ee Geenereeeee eee ee eeer eee re ee Per er ree eee eneeEee 174
FELIX sMARGIO— See MEIDAWANT (GABRIB Es ono oy 6. eee 156
FITZGERALD: S ——See: MARSHAITELE. SiAg 2 i5--msmer nse ae eer ener cee ee eee eee ene eee eee 641
FITZGERALD, SCOTT J.—Additional notes on Nearctic Bibio Geoffroy (Diptera: Bibionidae)
Pas MRR ociave oti teiebs Lie Be stereo lola wigedie-oletete 8 aantta nti cleleman rare tore atl nra cis er aerars nye hie Ree ee Cece ape eee 294
FITZGERALD, SCOTT J. and BORIS C. KONDRATIEFF—A new species of Pseudonomo-
aria eq unent (Dyes MbyelebrS)) timo IMIEXSEO) orossococgnosnaeconacencosdoobncosnnsosocooannbee 171
FOOTE WBIA:— See GARSON) IUEN) g2.505 ae sstenasceneceniee tieniaret eae nee eee meee ee ee ae eRe eee 541
FORATTINI, OSWLADO PAULO—See WILKERSON, RICHARD C. .....................00005 461
PULEERTON, STUART Mi See EVANS HOWARD IES eaeseeeeeene eee eer eee eee eee 174
GAGNE, RAYMOND J., JOSEPH K. BALCIUNAS, and DAMIEN W. BURROWS.—Six new
species of gall midges (Diptera: Cecidomyiidae) from Melaleuca (Murtaceae) in Australia
aiphaiale pib.ovtistaraioavain prenaninarefS.a Debuts eee ee a lane vate Raa ise a Mee aR rene A ne tere eee 312
GAGNE, RAYMOND J., CORNELIA OTT, and SUSANNE S. RENNER—A new species of
gall midge (Diptera: Cecidomyiidae) from Ecuador associated with flowers of Clavija (Theo-
PHEASCACEAE)! "Ff s ek ee beet eet eer ee oleae ot ON RR ett Bs SA nn 110
GANDOLFO, DANIEL E. and ALLEN L. NORRBOM—A new species of Trypanarestra
Hering (Diptera: Tephritidae) from Patagonia, a potential agent for biological control of
snakeweedsi(Gutierreziasspp» minsthes WiniteGins tate sine Eee eee eee ere eee enna 248
GOEDEN, RICHARD D. and JEFFREY A. TEERINK—Notes on the life histories and de-
scriptions of adults and immature stages of Procecidochares kristineae and P. lisae new
species (Diptera: Tephritidae) on Ambrosia spp. in southern California ........................ 67
GOEDEN, RICHARD D. and JEFFREY A. TEERINK—Life history and description of im-
mature stages of Procecidochares anthracina (Doane) (Diptera: Tephritidae) on Solidago
californica, Nuttallfingsouthernn Calitoniauees tree re eee ee eee Cee eeeee eeeee eee nee 180
GOEDEN, RICHARD D. and JEFFREY A. TEERINK—Life history and description of im-
mature stages of Xenochaeta dichromata Snow (Diptera: Tephritidae) on Hieracium albiflo-
jum Waookermunicentralyandesouthern) Calitonmauee-eeeeeeeeeeeee renee eee en eee eee eee eee ere eee 597
GOEDEN, RICHARD D. and JEFFREY A. TEERINK—Life history and description of im-
mature stages of Trupanea signata Foote (Diptera: Tephritidae) on Gnaphalium luteo-album
L. inSouthem: Califonntaon.i cere YS. tis ee yg et. eR ES pce eee 748
GOEDENWRIGHARDID Scab ADRICKS DAVID) En naee ate eee eee eae 415
COEDEN RICHARD ID See IBADRIGKe DAY ID ESIEe eather eee Eee eer pease eres 487
GRAB) Gi See: PIKE) KS. icine sep oe Soa ee OR Ee Ne ne en a ee eae 143
GRISSELL, E. E.—Biological notes on Sparasion Latreille (Hymenotera: Scelionidae), an egg
parasitoid of Atlanticus gibbosus Scudder (Orthoptera: Tettigonidae) ...................000..00s 693
VOLUME 99, NUMBER 4 781
HARRIS, STEVEN C.—See MOULTON, STEPHEN R., II
HEADRICK, DAVID—See SWEARINGEN, MICHEL
HEADRICK, DAVID H. and RICHARD D. GOEDEN—Gall midge forms galls on furit fly
callsi (Diptera Cecidomymidaemlephuitidac) yen.s pen: aeeeres ee eee aese cece acetate reereeecce ae 487
HEADRICK, DAVID H., RICHARD D. GOEDEN, and JEFFREY A. TEERINK—Taxonomy
of Aciurina trixa Curran (Diptera: Tephritidae) and its life history on Chrysothamnus nau-
seosus (Pallas) Britton in southern California; with notes on A. bigeloviae (Cockerell)
HERNANDEZ-ORTIZ, VICENTE—See MARTINEZ, IMELDA M. .................2.0c0eeee ee 127
HODGES, RONALD W.—A new agonoxenine moth damaging Araucaria araucana needles in
western Argentina and notes on the Neotropical agonoxenine fauna (Lepidoptera: Gelechioi-
Cea SE TACKS Tac) eee eee areata e cohdie SAS aleva dieisleceies bale Sale peas on ee Ae 267
HOEBE KEE SRICHARD——S cea WiHEb EE RewAtGawRa ceeasae ttn ete eat eerie nes a eee seer 409
WR Diese — Sect DUSSE. pRoAMEMOE BIN con ay ene nia ayh oe < sd Madirdsadd jonas Os oadhd woehdedae meee 490
JASCHHOE MATHIAS—Descriptions of a new genus and six new species of Nearctic Les-
tremiimac (Dipterass © ecid onmysicl ac) eee se meters tie arate mee Valetereers erste (eles areal a). oblate eth ele teil 523
JONES, ROBERT W. and HORACE R. BURKE—New species and host plants of the Anthon-
omus grandis species group (Coleoptera: Curculionidae) ................. 0... c cece eee eee 705
KANTO. K —See MURGEON Usk cnt cees seers ee ele oie ovcren) oe NioieieTala as aes eisai errsiele aietevster steers aie 608
KIRCHNER, RALPH E and BORIS C. KONDRATIEFF—A new species of Nearctic Perlesta
(Plecopteras Penlidae) irom Vanpimiayereeee sere esa ce acetate cee ee cier ec are rye ae ee iociar 290
KONDRATIEEEGBORIS| © See bIRAGERALD SS C Olga een emest ses ersseasee reece eer 171
KONDRATIEEEIBORIS €——Sce KIRCHNER SRAIER Hg Rey aeeee een eee reer reece eer eee ee 290
KOTRBA, MARION—Shoot or stab? Morphological evidence on the unresolved oviposition
technique in Stylogaster Macquart (Diptera: Conopidae), including discussion of behavioral
OL EAVEIMKOI NS) gapaououdsédoserodansesnoapcdoanosbobossh obo ockabancoconosoensoGt occ dpadomsucuaudccepecougne 614
KROMBEIN, KARL V. and BETH B. NORDEN—Nesting behavior of Krombeinictus norden-
ae Leclercq, a sphecid wasp with vegetarian larvae (Hymenoptera: Sphecidae: Crabroninae)
PCs ates OEE REO oe Deo er CU OCRR AES COaSO RO oct ee eon eter opr 5 Geer an ese ataae aan a Sunnnr 42
LARSON, LEN and B.A. FOOTE—Biology of four species of Notiphila Fallén (Diptera:
Ephydridae) associated with the yellow water lily, Nuphar luteum (Nymphaeaceae) ......... 541
BAND RYSBERNARD—See ADAMSIKi-D AWD) Meeps kee pekin state eeeceree ere ere eeceriacrt 348
LOYA, LANE J. and J.E. MCPHERSON—Life history and laboratory rearing of Oedancala
dorsalis (Say) (Heteroptera: Lygaeidae), with descriptions of immature stages ................ 89
LUGO-ORTIZ, C.R. and W.P,. MCCAFFERTY—A new genus and redescriptions for African
species previously placed in Acentrella (Ephemeroptera: Baetidae) ...............--- 2.60000 e ees 429
MACKAY, WILLIAM P—A revision of the Neotropcial ants of the genus Camponotus, sub-
genus Myrmostenus. (Hymenoptera: Formicidae) 2525.22.20... + saeco seis o> 22-2 = 194
MARSHALL, S.A.—Limomyza, a new genus of primitive Limosininae (Diptera: Sphaeroceri-
dae), with five new species from United States, Mexico, and Central America ............... 279
MARSHALL, S.A.—A first record of Aptilotus Mik (Diptera: Sphaeroceridae) from the Neo-
tropical Region, with the description of three new wingless species of the Aptilotus paradoxus
EACOLUYO) TaRoyoA lorvelol SMENAMNONNS 1 (COMED IRWES .scnaccoaceanecoqaaosnsan conn aancanno oso ve cagbdnGooncdeuc 505
MARSHALL, S.A. and S. FITZGERALD—Rudolfina cavernicola, a new species of cave-
associated Sphaeroceridae (Diptera) from Colorado and Arizona .................+---00seee eee 641
MARTINEZ, IMELDA M. and VICENTE HERNANDEZ-ORTIZ—Anatomy of the reproduc-
tive system in six Anastrepha species and comments regarding their terminology in Tephrit-
Gk: (WDM Gidea goegnctqseadosecGodomaonsdooodadidaaudcacepess cas dee toanos.dun aan abooonsdoDpEubUa Dt onoraseoror 127
IMUNIDEIENWS IDYAN INE See (CLOINI DOIN, IM UANIRIINC 5 caccoscocandoonadacnoveussacovopecer sbeoanaDuuDgaacccC 676
MATILE, LOIC—Fenderomyia Shaw, a valid North American taxon in Macrocerinae (Diptera:
My cetophiloideasNeroplatidac) meeerareematre ee eee ne tee eee eee eerie ias te ee ee 50
MATHIS, WAYNE N.—A review of the shore-fly genus Diphuia Cresson (Diptera: Ephydridae)
abiatiqn sagCR need ia Soe aa oda aiadaotagcudta accion Ga aanne ae aeureen can babe se naCaerincenacaneactnaceDadadr 28
MATHIS, WAYNE N.—A revision of Neotropical Ditricophora Cresson (Diptera: Ephydridae)
Bo Cg PEGE SUR er icc Wane not tnaG Conn At acd Gina ane cape 7oG Dan De (OUOmrnre ema Anan son Gotan qoabeasooerbe 697
INK C/N AIEIBRMENG WAP — See ILLUKGOROIRINUZ, (CAR, cooccncde nnasb enpoansodohnes0b|o0s00 905000090 000G ORE 429
IMIGPEIA RON, Va See IOC ILZNINIE) Ul, cacacaseanscsden0000000 ccnp a aceneacsHb0 seo ooaU DROS ATO BOeE 89
MEJDALANI, GABRIEL and MARCIO FELIX—A new species of the Neotropical genus
Lissoscarta Stal (Homoptera: Cicadellidae: Cicadellinae) that mimics wasps .......-.-.--..-+- 156
782 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
MELIKA, GEORGE and WARREN G. ABRAHAMSON—Descriptions of four new species
of cynipid gall wasps of the genus Neuroterus Hartig (Hymenoptera: Cynipidae) with redes-
criptions of some known species from the eastern United States ......2...22.2...-.2-..-4-2---- 560
MELIKA, GEORGE and WARREN G. ABRAHAMSON—Synonymy of two genera (Eumay-
ria and Trisoleniella) of cynipid gall wasps and description of a new genus, Eumayriella
(Hymenoptera? (Cyniprdae)" % 5252s G sete oes ei ee ae Ie etait s ole fia Disfave nbeloes Ghcters cretereToietovers tee eee 666
MENKE, ARNOLD S.—A new Ammoplanus Gussakovskij (Hymenoptera: Sphecidae: Pem-
phredoninae) tromySaniC@lementedslandsCalitonailasereene:-ceee eas ee eee ee eee eee ee eee 183
MIER DURANTE, PILAR M., JAIME ORTEGO, and JUAN M. NIETO NAFRIA— A new
subgenus and species of the genus Brachyunguis Das (Hemiptera: Aphididae) from Argentina
Be ec So te ear c aCnn nace con socio nua aETa tr eM oerrctc inca tocar seAcoach tad daqhour 720
MILLER, DOUGLASS R. and SRIDHAR POLAVARAPU—A new species of mealybug in
the genus Dysmicoccus (Hemiptera: Coccoidea: Pseudococcidae) of importance in highbush
blueberries (Vaccinium corymbosum, Ericaceae) in the eastern United States ................. 440
MILLER, DOUGLASS R. and DOUGLAS J. WILLIAMS—A new species of mealybug in the
genus Pseudococcus (Homoptera: Pseudococcidae) of quarantine importance ................. 305
MILLER, KELLY B.— Crinodessus amyae, a new Nearctic genus and species of predaceous
diving beetle (Coleoptera: Dytiscidae: Hydroporinae: Bidessini) from Texas, U.S.A. ......... 483
MILLERS T==Sée PIKES (KS )oh 2s 5 dss net eo ele ois a ere RSE nee eee Re ee ean 143
MORRONE; JUAN J.—-See‘COSCARON) MARTA DELiGC. : 5:25) ee ee 55
MOULTON, STEPHEN R., II and STEVEN C. HARRIS—New species of southwestern Ne-
arctic muicrocaddistives 1 @nichopteta hly droptilidae) memsnse-eeere eter eee eee eeaer cee eee seers 494
MUNOZ-QUESADA, FERNANDO—Five new species and a new record of Costa Rican Lep-
tonemas Guerin @irichoptera Ey dropsy chidde)meeesecerereessteee eee arene cee eeeee tere eee ee ene eeee Ls;
NIETO NAFRIA, JUAN M.—See MIER DURANTE, PILAR M. ............0.0.0.0c0c0eeee eens 720
NORDEN} BETH B:—See INROMBEINARARID Va ane tecacee pect enna eae eee 42
NORRBOMPALLEN IE See GANDOEROF DANTE TB aa nee et eee ee eee eee eee eee er ee 248
NORRBOM, ALLEN L.—The genus Carpomya Costa (Diptera: Tephritidae): new synonymy,
description of first American species, and phylogenetic analysis .....................000eeee000: 338
ORDEGOSJAIME——ScesMIER DURANTE PIVARIM Seceseee eect ee eee eee a eee er eee eer ee ree 720
OTR iCORNELIA—SceiGAGNE “RAYMOND ites, eee ee ee eee eee 110
PIKE, K.S., P. STARY, D. ALLISON, G. GRAF L. BOYDSTON, and T. MILLER— Parasitoids
(Hymenoptera:Braconidae: Aphidinae) of aphids on big sagebrush (Artemisia tridentata Nut-
tall) and prairie sage (Artemisia ludoviciana Nuttall) in Washington State ..................... 143
PINTO, JOHN D.—Trichogrammatoidea brasiliensis (Ashmead)—new combination for a spe-
cies historically placed in Trichogramma (Hymenoptera: Trichogrammatidae) ................ 595
PINTO, JOHN D., RICHARD STOUTHAMER, and GARY R. PLATNER—A new cryptic
species of Trichogramma (Hymenoptera: Trichogrammatidae) from the Mojave Desert of
California as determined by morphological, reproductive and molecular data ................. 238
PLATNER; ‘GARY R:=—See PINTO: JOBING iio. eee eee eer nee ere See eee 238
POINARS GEORGE? OMIR— See ZUPARKOSROBE RTD pee eee ester een eee eer eE Eee 744
POLAVARAPUSSRIDHAR—=_See MIEEBR] DOUGEASS IR eset eee erent eee Rees 440
PUN; WING-WAH=See EASTON? EMME TR a areedeconin cs aetee aeehise emer nie eee eene Ree ee 574
RADOVIC, IVICA and SRDJAN SUSIC—Morphological characteristics of the sting and prey
carriage mechanism in Sericophorus relucens F Smith (Hymenoptera: Sphecidae: Larrinae)
wistaieleiclats MARA At ROA pea i dette tee teresa aerate ctane velo tra clase wate a TOISTO Se cect at Fayence ete rt inte mo eT sya eee 537
RENNER; SUSANNE: S:——SeeGAGNE; RAYMOND J. ...c.sishncsteas see aeanee eae enone 110
ROBINSON, HAROLD and MARK DEYRUP—Two new species of Asyndetus Loew, and
notes on the identity of A. interruptus Loew (Diptera: Dolochopodidae) .....................-. 477
ROSE, MIKE and GREGORY ZOLNEROWICH—Eretmocerus Haldeman (Hymenoptera:
Aphelinidae) in the United States, with descriptions of new species attacking Bemisia (tabaci
complex) (Homoptera: Aleyrodidac)teas.2---eeee ese e ee eee eee ee eee eee eee eee ee tee ee eee 1
SALLUM, MARIA ANICE M.—See WILKERSON, RICHARD C. ..............20eeceeeeeeeeees 461
SANTIAGO-FRAGOSO, SILVIA—See BUENO-SORIA, JOAQUIN ...............000cceeneeenes 859
SCHAEFER, CARL W. and JOSEPH C. SCHAFFNER—The New World alydine genus Stach-
yocnemmus stal (Hemiptera: Alydidaé)) 2 .:..nsccscneet cohen ee boon oe aes eee 136
SCHAEENER- JOSEPH €——SeetS CHABFER CARTS Wit ceaeteeerernete a eeeeee ee ee eee 136
SCHAUFR M.E.-—See WHESERA RAV Gea We ec ee ee EIT AE eee eee ee 101
VOLUME 99, NUMBER 4
SCHIBEBRAGERENCEWe-——SeerS MIME aD A VID BRS (sma -meccisemctectncecetn densa cals eceeiaeee
SLATER, JAMES A. and HARRY BRAILOVSKY—A new species of Drymus Fieber from
Mexico, with a key to species and a checklist of Western Hemisphere Drymini (Hemiptera:
WE ACTA AS) ee ete eee SI cet CLS oos oie ccaiSinresbsasdls wine suds Bae BemAGE SAGE odie Boxee de ohare eee
SMITH, DAVID R.—Aulacidae (Hymenoptera) of Sri Lanka ...........00..0 0 ce cece cece eee eeeee
SMITH, DAVID R. and TERENCE L. SCHIEFER—A new genus and species of Allantinae
(Hymenoptera: Tenthredinidae) from southeastern United States ......................0222.. 20
SUN RNG TR Sas ERIS LOS cc spn ae aose 08. na ie ee ree
STOUMTAMER] RIGEARD—— See RINMOAIORINGD ie. seo ce scacce sc ceiee eee ceee ee ceeeeeeeetene
SUISECL SIDIVNIS Sa RYAN DION/ CL INC ae hd en Oe: oo eee
SWEARINGEN, MICHEL, DAVID HEADRICK, and TOM BELLOWS— Comparison of fix-
ation and drying procedures for scanning electron microscopy among insect body types ....
TEERINK JBEEREWAj——SeerGOEDEN, RIGHARDID) 2 iosceeecceec soc -ckece cc. se seneseeedeeeee
TEV UIN KS US SI RIENG ZN Sas) (G18) D) INL, RI KCIGUANRID) ID), Sasancnonosonsnaanquondecds sadcoconpeseesn
MEBRINKVERBREYeAC—SseelGOEDENSRICEHAR DID Were scea eee sseeeteaseaeecee ce ee eeeen ce:
THEIRINIK, USAR A= See GOURD BING IRIKEISUNIRID) ID), se dcnpcs0c0ncn sbocs0euceocbdousnasnabnanoe
TBS RUIN, MSR ARI EN. A — See Isl EVADINICCIS IDVANID) 1S saacoooscoonombaacganocssbooduoovnsesoconne
TENNESSEN, K. J.— Lestes jerrelli, n. sp. (Zygoptera: Lestidae), a new damselfly from Ec-
(EVAL P seomtn perce de ch Ob ee ateracn BER EA rrcn Roan serie crars Reo sates oman ORR Ia tein) eG APOE Ir PORT eC CeCe TSE
THOMPSON, FE CHRISTIAN—Revision of the Eristalis flower flies (Diptera: Syrphidae) of
therAinenicasssouthvotsthe MWmiteds S tate sacar ayaa eiareterstetsrar tars sratetel overs te vevelerctete siete clare aera
THOMPSON, E CHRISTIAN—The genus Pleskeola Stackelberg (Diptera: Syrphidae), a junior
SyNOMY iol Ole Lar nelopnilus Girschnetesseceeee-ceeeere sees ae eeeeeeeeeecscce see oeisiseseeee a
TOGASHI, ICHIJI—A new genus and species belonging to the subfamily Sterictiphorinae
(Hymenoptera; SymphytaeAgsidac) miromapaneerescree se eeeeerere cesses sea ceseteisecene
TOGASHI, ICHIJI—A new genus and a new species belonging to the subfamily Blennocam-
pinaer(Elymenopteraswenthredinidac) irom apanwee seme eric eseeseeeeeasie tesieeee eer
TRUEE OVE ANN—See CONDONE WEAR YG ae ceretessese aisles sie safslclols Sieisisislofai tes eletec exe slates s nvsicistorsteuslaiete cies
TURGEON, J. J.. K. KAMIJO, and G. DEBARR—A new species of Megastigmus Dalman
(Hymenoptera: Torymidae) reared from seeds of Atlantic white cedar (Cupressaceae), with
MNOtesvOninfestatiOnyratesh ae drsetsee aaa cia seeders eels tine cael stay ae enon Ee ee Strela eaves
WHEELER, A.G., JR.—Empoasca (Kybos) luda Davidson and DeLong: distribution and habits
of an immigrant birch-feeding leafhopper in North America (Homoptera: Cicadellidae) .....
WHEELER, A.G. JR. and E. RICHARD HOEBEKE—YTrioza chenopodii Reuter: first North
American record of an Old World psyllid (Homoptera: Psylloidea: Triozidae) ................
WIJESEKARA, G.A.W. and M.E. SCHAUFF—Twq new genera and three new species of
Buplectnnin(iymenoptera:; sulophidae) arom the yINew World ere. on-screen eee ne
WILKERSON, RICHARD C., MARIA ANICE M. SALLUM, and OSWALDO PAULO FOR-
ATTINI—Redescription of Anopheles (Anopheles) shannoni Davis; a member of the Arri-
balzagia series from the Amazon Basin (Diptera: Culicidae) ...................... 0. eee e eee eee es
WIEEIAMS DOUGLAS J——Sce MILE BReDOUGIEASS! Raeraeneees renee reisecreceeasterecir
ZAOUNEROWICHE GREG ORYA—ScesR@SEss MIKE Serre ee eaaee ere ee eecceaeeee een err: cece
ZUPARKO, ROBERT L. and GEORGE O. POINAR, JR.— Aivalykus dominicanus (Hyme-
noptera: Braconidae), a new species from Dominican amber ..................-.2+-eeeeeeeeeeees
NOTES
AINAGNOSEAKISS SAND WAGE = Seer PAIKCATE UKE RAIMI Seer eee cert elects elcleres aletete
DUFFIELD, RICHARD M. and CHARLES H. NELSON—Note on stoneflies (Plecoptera),
particularly Prostoia besametsa (Ricker) (Nemouridae), in the diet of salmonids from the
headwaters of the middle fork of the South Platte River, Colorado ....................0..0.00-+
EPSTEIN, MARC E.—Parasa indetermina (Boisduval) (Lepidoptera: Limacodidae), a new host
Loney StOpUsSimacer loew, (Dipteras bombyliidae) meres se seEee eee eects ees sis ieee
FAGERLUND, RICHARD—See MACKAY, WILLIAMP. .................-..e cesses eee eeteeeeees
GAGNE, RAYMOND J. and JOHN C. MOSER—Celticecis, a genus of gall midges (Diptera:
Cecidomyiidae), newly reported for the western Palearctic Region .....................20225---
MACKAY, WILLIAM P. and RICHARD FAGERLUND—Range expansion of the red imported
fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae), into New Mexico and extreme
WIGS NORA coo addsdeoss souscondocecuupgedas cone te reEE er onaea Rood OCt Uae Uap ac cT SC CCe Eo ene eT iar ia
472
661
608
623
409
101
784 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
MARSH, PAUL M.—Replacement names for Western Hemisphere genera of Doryctinae (Hy-
menoptéera: Braconidae) 322: esas oh at oda cemomnee ee Eee ete a clea ee ee ete eee ee eee 586
MOSER; JOHN 'C.—See 'GAGNE: RAYMOND). eis oe eee ee 756
NELSON, .GHAREES H-—Sce DUBEBIBIDS RICHARDS aaeeeer eee tee - ernee epee eee aeeeee 374
MENKE, ARNOLD S.—Species homonymy in Belomicrus Costa (Hymenoptera: Sphecidae:
Crabroninae))é «,. sii J ieladv ascends AAD Woe, ce eh op reels eae PET RE PEEP ROAR Ceo etotetol ce ae one nee 378
PAKALUK, JAMES and SANDRA L. ANAGNOSTAKIS—Acoptus suturalis LeConte (Co-
leoptera: Curculionidae: Zygopinae), a potential vector of the chestnut blight fungus, Cry-
phonectria parasitica (NMurnll) Barr anwtheyeastem United: Statese-nesep eee eneee ee eee nee 583
PALMA, RICARDO) L..—See:- PRICE ROGERYD gree ceee eee eae eee eee Cree eee eee ee eee 587
PRICE, ROGER D. and RICARDO L. PALMA—Gender correction for a louse (Phthiraptera:
Menoponidae)' from. Cormorants i i3 pene en riean retire tora aio ace cte See oe eee eee 587
SCHAEFER, CARL W.—Brachycerocoris camelus Costa (Hemiptera: Pentatomidae: Podopi-
nae) in India: newscountry TECOrd) so. 54.552 5- hams eee naire ee Oe oe CE OE ee 204
SMITH, DAVID R.—Collections of Stephanidae (Hymenoptera) in the mid-Atlantic states,
including an eastern record for Schlettererius cinctipes (Cresson) .............+.00eeeeeeeeeeeees B/T/
STAINES, C. L.—Cereal leaf beetle (Coleoptera: Chrysomelidae) as a pest of ornamental
QTASSES™ hp achese a terae sae eed o/h nie sl Sis nnn Siale nhs scevarwle cle aIeIE Ae reye elo Gicrapeley ltrs ere ere oretepere otstere state ae teeraiers eee 376
Book REVIEWS
BRODA-HYDORN, SUSAN— Thrips Biology and Management, edited by Bruce L. Parker,
Margaret Skinner ‘and Trevor Eewis -a--cccssceesce coerce er oe nent eee eee eee eee eae 205
HORMIGA, GUSTAVO—Riceland Spiders of South and Southeast Asia, by A. T. Barrion and
AL ETSI en’ she te aaa sos erie tlecsing oo cmeactoa ae ore «Hees apa ek i a ee 399
MILLER, GARY L.—Brethren of the Net: American Entomology, 1840-1880, by W. Conner
S10]c2 112 | en aon een ie Panels MARAE ear Ad onriae Goctise das cchon cS56be Adc rlaadorénooeth oe 758
OBITUARIES
BCKEREIN, RALPH P— See ROBBINS RIGHA RID iIGaee secrete eee eee eee eee eee Eeaeee 588
HENRY THONAS I ——Sceis ANITMAGO=BIPAY. TOR GE At ape ren teeeeere ee eee ene eee 762
ROBBINS, RICHARD G. and RALPH P. ECKERLIN—Robert Traub (1916-1996): an appre-
ei: 1010) US aESpr CEMA an reer an He AS Sat AeOROR Se AE CAPM aca at sae auocatany Soh cane sb DALADHas oon as oeaacoona: 588
SABROSKYA CURTIS MW-— George) G7 steyskali(i909= 11996) pia a eee ee 379
SANTIAGO-BLAY, JORGE A., THOMAS J. HENRY, and A.G. WHEELER, JR.—Jenaro Mal-
donado Capriles (9191995) eS ysiere aeae A se ets ce ee Saar Ee 762
WHEELERGAG.IR-——See SANTMIAGOABIEAYS JORGE WAG eae here Cee EEE ere 762
MISCELLANEOUS
INSTRUCTIONS RORVAWITHORS. Beasts are See eC Soe oe eee eee 591
OFFICIAL SEAL OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON .............. 208
REPORTS; OPOBBICER SS) hae sce ct eo terae slasGi nae kOe ee oe EERE: OORT Eee REECE eee eee 402
SOGITE TYGIMIB ERIN GS 3 we srewaspin see secs nlehe Giada ace EELS Beco EE eee 404, 772
VOLUME INDIE Ket oc setatertiocs bast oe cre tele a ara vies, ge see eee aces ee 778
PUBLICATIONS FOR SALE BY THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON
MISCELLANEOUS PUBLICATIONS
Cynipid Galls of the Eastern United States, by Lewis H. Weld. 124 pp. 1959
Cynipid Galls of the Southwest, by Lewis H. Weld. 35 pp. 1960
Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C. Steyskal.
35 pp. 1977 En
Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C.
SS Tey SRA COL OY 7 pana a SR oe 8 SE ee Nk
A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael E.
Schauff. 85 pp. 1990
A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera): Second Edition, Revised, by E. Eric
SrissellcandentichaeliS: Schatitta soins i yc emeate Sek eee 0 ss a, Se Me
MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 —
No.2. A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving.
95 pp. 1942 _— ua Bo 0S. BFC: fe
No.3. The Nearctic Leute. a Generic Classification and Check List, by Paul Wilson Oman. 253
pp. 1949 - Bee IE SA SIS So MN BAI 8 LA Re 2 La era SO me ae BAe ee
No. 4. A Manual of the Ae by G. W. Wharton and H. S. Fuller. 185 pp. 1952...»
No.5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 ___
No.6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi
Takahasi. 230 pp. 1969
No.7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp.
1976
No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato-
poponidae), by W.)L: Grogan, Jr, and:“W..W.. :Wirth/ 125. pp. 1979.2 ee
No.9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson. 200 pp.
1981
No. 10. Recent Advances in Dipteran Systematics: Commemorative Volume in Honor of Curtis W.
Sabrosky. Edited by Wayne N. Mathis and F. Christian Thompson. 227 pp. 1982
No. 11. A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370 pp.
TBS ee Were te or eye eee tae tent 2 ANE) Ree Sve ah noe BES DL Ae Ae tk ee Te
No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff.
67 pp. 1984
No. 13. An Identification Manual for the North American Genera of the Family Braconidae (Hyme-
noptera), by Paul M. Marsh, Scott R. Shaw, and Robert A. Wharton. 98 pp. 1987 __-
No. 14. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174
pp. 1995
No. 15. A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent
arya en Grocan. Jemima me diay take cera iia Ne ee ee ot ee
No. 16. The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 —
No. 17. Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Krombein, edited
Dyib wb. NordentdnduATS. Melke-21o;ppilO96 tse Se eet 2 ee ee eS
No. 18. Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and
William L. Grogan, Jr. 297 pp. 1997
No. 19. Monograph of the Stilt Bugs, or Berytidae (Heteroptera), of the Western Hemisphere, by
Tlavepagre ise ee 8 Ce) ave E'S) yoyo) IRON p ea tay Cae A ee ee ee eee
5.00
18.00
25.00
25.00
25.00
25.00
25.00
18.00
Back issues of the Proceedings of the Entomological Society of Washington are available at $60.00 per volume
to non-members and $25.00 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, % Department of Entomology, MRC 168, Smithsonian
Institution, Washington, D.C. 20560.
CONTENTS
(Continued from front cover)
MIER DURANTE, M. PILAR, JAIME ORTEGO and JUAN M. NIETO NAFRIA—A new
subgenus and species of the genus Brachyunguis Das (Hemiptera: Aphididae) from Argen-
MELIKA, GEORGE and WARREN G. ABRAHAMSON-—Synonymy of two genera (Eumayria
and Trisoleniella) of cynipid gall wasps and description of a new genus, Eumayriella (Hy-
menoptera: Cynipidae)
PINTO, JOHN D.—Trichogrammatoidea brasiliensis (Ashmead)—new combination for a spe-
cies historically placed in Trichogramma (Hymenoptera: Trichogrammatidae)
TENNESSEN, K. J.—Lestes jerrelli, n. sp. (Zygoptera: Lestidae), a new damselfly from Ec-
THOMPSON, F. CHRISTIAN—The genus Pleskeola Stackelberg (Diptera: Syrphidae), a junior
synonym of Parhelophilus Girschner
TURGEON, J. J., K. KAMIJO and G. DEBARR—A new species of Megastigmus Dalman (Hy-
menoptera: Torymidae) reared from seeds of Atlantic white cedar (Cupressaceae), with notes
on infestation rates
WHEELER, A. G., JR.—Empoasca (Kybos) luda Davidson and DeLong: distribution and habits
of an immigrant birch-feeding leafhopper in North America (Homoptera: Cicadellidae) ....
ZUPARKO, ROBERT L. and GEORGE O. POINAR, Jr.—Aivalykus dominicanus (Hymenop-
tera: Braconidae), a new species from Dominican amber
NOTES
GAGNE, RAYMOND J. and JOHN C. MOSER—Celticecis, a genus of gall midges (Diptera:
Cecidomyiidae), newly reported for the western Palearctic Region
MacKAY, WILLIAM P. and RICHARD FAGERLUND—Range expansion of the red im-
ported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae), into New Mexico and
extreme western Texas
BOOK REVIEW
MILLER, GARY L.—Brethren of the Net: American Entomology, 1840-1880, by W. Conner
Sorensen
OBITUARY
SANTIAGO-BLAY, JORGE A., THOMAS J. HENRY and A. G. WHEELER, Jr.—Jenaro
Maldonado Capriles (1919-1995)
MISCELLANEOUS
Society Meetings
Volume Index
666
593
661
628
758
Vee
\ AT
py i 4 '
“ Daag yi"
day , i |
pra , oie \
' re j
iY
a
meee
"sh
ve i,
j j
Wek eine
ve a gS
said o fie
AW
MA i
HECKMAN lal
BINDERY INC. =
FEB 98
Set =~ Tea RE PART (OC CL Ete
(eng aky “
Cea an Tae
WB NCA en
Deer wr
Nea
PM PAARL AEM ME UAE LE
es ee ae
DAA MERE Oe
eA: MONCH. KER
PAE DUA APL
V2 adie bea ebte ba Nal
eer er uy re ent
Wor MeL a) hab al
ea
Bi hie Mh CA lo GR
CHEM By
’ P) ay ’ a3 ' i
. : Seige, " \ ri ee 4 x
\ ufig \
HN ifthe 7 \ . . ‘
Uae 7 Mie 4 bc an to \ «
Pon mn UA ihe ae i
Me ‘ ‘ fl ~ " au r .
mee a ; M Bye May ; { :
MUS UAG . ‘ ‘ : ‘ , ‘ 4 ;
bt 1 eth aes WS Sh RL vitehadtedees asi tyesb era gen uted ree ete ay ‘ wow t * f wi
Live Music Archive
Librivox Free Audio
Metropolitan Museum
Cleveland Museum of Art
Internet Arcade
Console Living Room
Books to Borrow
Open Library
TV News
Understanding 9/11