ares 


Z hens aratt arene 
. oe Whe la dip he a a plete te elatad Mets Pha hagas atin tl ity Vivek phate betssa bet 
Pre Nef wel a BEE hie ba ae ee Le beb eaten 


Viegegiyt et 
eee as 
vente 

PV bane 


Meant VPI Geek 
feccen Pinas 


Soh a 


Fa dB Be hat walt 


mR pte ade 


Ba NEATE oN 
Saeabatie uoenenrr 


eh ete te 
EAE Se 
Vepee ake 
eerey 
ererer ent 
apress 


Teh EL 


RiONunese 


eee rays 
Trae erener! 


Ba ter cteuny 


Sy 


Mah ateewtahane 


RPP dated 
Pet ta a tee 


vet 
ao 


lated ghar ene 
EVV EIEN at tg 


ke 

ala 

to JULY 2000 NO. 3 
faa ca Gb. (ISSN 0013-8797) 


Tat 


“PROCEEDINGS 


of the 


ENTOMOLOGICAL SOCIETY 


VOL;; 102 


CONTENTS 


BAPTISTA, ALESSANDRA R. P. and WAYNE N. MATHIS—Notes on the genus Cyamops 


Melander (Diptera: Periscelididae), including description Of fen NEWASPECIES! Phebe eyo: eiaiele 481 
BRASWELL, W. EVAN and JAMES R. OTT—The biology of Doa ampla (Grote) (Lepidoptera: 
Doidae) on its host plant Stillingia texana (Euphorbiaceae) .......---...++-+ see eeeee serene 507 


BROWER, ANDREW V. Z.—On the validity of Heliconius tristero Brower and Heliconius 
melpomene mocoa Brower, with notes on species concepts in Heliconius Kluk (Lepidoptera: 
INivaniphalidae) ie. shes oa spats ae Sew send d sieved oe a plv ne a4 tele es piineg een rieueee ones aaah. 678 


DARSIE, RICHARD F., JR.—The pupae of three species in Minomyia Theobald from Nepal and 
a key to the known pupae of the genus in the Oriental and Australasian regions (Diptera: 
Sagat Pe, uiatsie cla 2 ayo oats hrs halalgnsee Pama tt oni k ale eo ma wenac ere «gence ans onniee ee neo 625 


FOSTER, GEORGE A. and WAYNE N. MATHIS—Notes on Neotropical species of Tethina 
Haliday (Diptera: Tethinidae) ........0.. 20... escent eee cet eee eden treet eb ces seen ee nec cn ne rcdes 542 


GAIMARI, STEPHEN D. and MICHAEL E. IRWIN—Revision of the mexicana-group of the 
cycloteline genus Ozodiceromyia Bigot (Diptera: Mherevidae) io. ota. asceaee eee eee er 561 


GOEDEN, RICHARD D.—Life history and description of immature stages of Neaspilota appen- 
diculata Freidberg and Mathis (Diptera: Tephritidae) on Machaeranthera canescens (Pursh) 


A. Gray (Asteraceae) in southern California .........--..0++ + sees sere eee ecr teeter tees tsees S19 
HASTRITER, MICHAEL W. and EUSTORGIO MENDEZ—A review of the flea genera Hectopsylla 
Frauenfeld and Rhynchopsyllus Haller (Siphonaptera: Pulicidae) .........-.-++-++-+0+eeres eee ees 613 
KRUSE, JAMES J.—Archips goyeriana, n. sp. (Lepidoptera: Tortricidae) an important pest of 
baldcypress (Taxodiaceae) in Louisiana and MUSBISSIDpi fee st sys eee aot See pes arlts eine sg 741 
LAPIERRE, LOUIS M.—Prey selection and diurnal activity of Holcocephala oculata (F.) (Diptera: 
Asitidae) 10 Costa Rica. ois 3.0... ee eee tes goemmn nd yasanie eres astitenar eb ser cna dele Ssma)is 643 


MAIER, CHRIS T. and CAROL R. LEMMON—Discovery of the small Japanese cedar long- 
horned beetle, Callidiellum rufipenne (Motschulsky) (Coleoptera: Cerambycidae), in live 
arborvitae in Connecticut 0. ....02.2 ccc e cece ete te cnc tele cere nec clne ce celence te dee sits cineceeecees 747 


MARSHALL, S. A.—Chespiritos, a new genus of Limosininae (Diptera: Sphaeroceridae) from 
GO STAMR TGA ete ae ed Uva neh estas Me Lics Pe Siare ere ctubate ata: cf Mars USI PRISI in elagare/s stelers lee ais ieteialejols eye Sia 609 


(Continued on back cover) 


THE 


ENTOMOLOGICAL SOCIETY 
OF WASHINGTON 


OFFICERS FOR 2000 


Davin G. Furtu, President MIcHAEL G. PoGug, Treasurer 
JOHN W. Brown, President-Elect THEODORE R. ScuHuLtz, Program Chair 
Stuart H. McKamey, Recording Secretary STEVEN W. LinGAFELTER, Membership Chair 
Ho tis B. WILLIAMS, Corresponding Secretary MiIcHaeEL E. Scuaurr, Past President 


Jon A. Lewis, Custodian 
Davip R. Situ, Editor 


Publications Committee 
RAYMOND J. GAGNE THOMAS J. HENRY Wayne N. MArTHIs 


Honorary President 
Louts—E M. RussELL 


Honorary Members 
Kart V. KROMBEIN RONALD W. HopGEs 
DONALD M. ANDERSON WILLIAM E. BickLEy 


All correspondence concerning Society business should be mailed to the appropriate officer at the following 
address: Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Wash- 
ington, D.C. 20560-0168. 


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 7:30 PM. 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, Smithsonian 
Institution, Washington, D.C. 20560-0168. Members in good standing receive the Proceedings of the Entomo- 
logical 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. 771-772 OF THIS 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. 20560-0168. 


Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, 
Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. 


Books for Review: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of 
Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. 


Managing Editor and Known Bondholders or other Security Holders: none. 
an ONES Lr 
This issue was mailed 14 July 2000 


Periodicals 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). 


v 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 481-506 


NOTES ON THE GENUS CYAMOPS MELANDER (DIPTERA: 
PERISCELIDIDAE), INCLUDING DESCRIPTION OF TEN NEW SPECIES 


ALESSANDRA R. P. BAPTISTA AND WAYNE N. MATHIS 


Department of Entomology, National Museum of Natural History, Smithsonian Insti- 
tution, Washington, D.C. 20560-0169, U.S.A. (e-mail: baptista@nmnh.si.edu and 
mathis.wayne @nmnh.si.edu) 


Abstract.—The species of Cyamops Melander are reviewed for five regional faunas, 
including the description of 10 new species as follows: C. funkae (Guyana), C. nigeriensis 
(Nigeria), C. freidbergi (Madagascar), C. micronesicus (Yap), C. fiji (Fiji), C. samoensis 
(American Samoa), C. femoratus (Philippine Islands), C. laos (Laos), C. banvaneue 
(Laos), C. kaplanae (Thailand). Cyamops pectinatus Khoo is reported from Tasmania. 
Regional keys are provided for the New World, Afrotropical, Australasian/Oceanian, and 
Oriental faunas. 


Key Words: Diptera, Periscelididae, Cyamops, New World, Afrotropical, Australasian/ 


Oceanian, Oriental, Palearctic 


The genus Cyamops Melander includes 
14 valid species: three in the Nearctic Re- 
gion, five in the Neotropical Region (Bap- 
tista and Mathis 1994, 1996), and six in the 
Australasian Region (Khoo 1985). Since re- 
vising the New World species of Cyamops 
(Baptista and Mathis 1994), we (Baptista 
and Mathis 1996) have described one ad- 
ditional species, C. sabroskyi, from speci- 
mens collected in Rio de Janeiro, Brazil. 
When we published our revision of New 
World species, several new species from lo- 
calities elsewhere in the world were then 
known to us, and some of these species had 
previously been reported (Hennig 1969, Sa- 
brosky 1980). The purpose of this paper is 
to describe most of the new species and up- 
date the existent keys to facilitate identifi- 
cation of all known species of Cyamops. 
The new species that we studied and that 
are not described in this paper are repre- 
sented by poorly preserved specimens, fre- 
quently a single male. 


Methods.—The descriptive terminology, 
with the exceptions noted in Baptista and 
Mathis (1994), is that published in the Man- 
ual of Nearctic Diptera (McAlpine 1981). 
As we recently published a complete de- 
scription for the genus Cyamops, the ge- 
neric characters are not repeated here. The 
format for species’ description likewise ad- 
heres to Baptista and Mathis (1994), with 
the following modifications: 

Facial shape: In addition to being sexu- 
ally dimorphic, two basic facial shapes oc- 
cur among males of Cyamops: 1. Face con- 
stricted medially by the anteroventral mar- 
gin of the eyes, expanding into a triangular 
region ventrally below the level of the pseu- 
dovibrissae and bearing a median ridge that 
was called the “facial carina” in our first 
paper. The corresponding female face is 
trapezoidal below the level of the pseudov- 
ibrissa, with a central, large, more elevated 
area. 2. Face not constricted medially, with- 
out a median ridge. The corresponding fe- 


482 


male face is equally large but has no cen- 
tral, elevated area. These shapes will be cit- 
ed simply as ‘“‘face of male narrowed” or 
‘not narrowed.” 

The structure we called the “‘hypandrial 
projection”’ in our revision will not be used 
in species descriptions herein, as this struc- 
ture is apparently unique to the following 
Neotropical species: C. halteratus Sabros- 
ky, C. nebulosus Melander, C. buenorum 
Baptista and Mathis, and C. faciatus Bap- 
tista and Mathis. This structure is men- 
tioned in the revised key, however, as it dis- 
tinguishes the Neotropical species just not- 
ed. 

The ‘“‘Key for the Australasian and 
Oceanian species of Cyamops”’ was gener- 
ated using DELTA (Dallwitz et al. 1998). 
The character states for the Australian spe- 
cies, described by Khoo (1985) and Hennig 
(1969), were taken from their publications. 

Species represented by poorly preserved 
specimens are noted and are included in an 
appropriate regional key. Our purpose in 
providing information on undescribed spe- 
cies is to call attention to character variation 
and distributional data for future studies on 
Cyamops. 

Although most specimens for this study 
are in the National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
D.C. (USNM), we also studied numerous 
specimens that were borrowed from and 
will be deposited in the Bernice P. Bishop 
Museum (BPBM). 


NEOTROPICAL REGION 


The new species treated in this section 
all key to C. americus Baptista and Mathis 
in our “‘Key to the New World species of 
Cyamops Melander”’ (Baptista and Mathis 
1994) and form a species group that is char- 
acterized by the following combination of 
characters: face of male constricted medi- 
ally; crossvein bm-cu present, separating 
cell bm from cell dc. To help identify the 
new American species discovered since our 
revision, we have furnished a revised key 
to the New World species. Species of the 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


genus Cyamops that have large geographic 
distributions, however, demonstrate some 
variation in the extent of microtomentum 
and coloration of the legs and wings. Thus, 
study of characters of the male terminalia 
is still advisable to determine a species’ 
identity accurately. 


KEY TO NEW WORLD SPECIES OF CYAMOPS 


1. Anepisternum bare along posterior margin ... 2 
— Anepisternum setose along posterior margin, 
usually bearing | seta and a few setulae ... 6 
2. Only apical scutellar seta present (United 
States) essere eee C. imitatus Sturtevant 
— Apical and basal scutellar setae present.... 3 


3. Wing hyaline (Canada, United States) ...... 

C. halteratus Sabrosky 

— Wing at least partially infuscate 

4. Vein R,,, sinuous; mid- and hindtibiae mostly 
yellow; scutellar disc slightly convex (eastern 
Canada and United States) ............... 


— Vein R,,,; straight or nearly so; mid- and hind- 
tibiae lightly infuscate to brown; scutellar disc 
flaty 3.38. c ha eens. 2 Beep ee eee eee 3 
5. Male right surstylus as narrow as left; hypan- 
drial projection large and exposed; 7th and 
8th sternites reduced (Brazil) 
C. fasciatus Baptista and Mathis 
— Male right surstylus much broader than left; 
hypandrial projection of normal length, not 
large, usually hidden beneath right surstylus; 
7th and 8th sternites well developed, fused to 
Sthistenite (iIVMextco) iene ae nen 
Rater CRC C. buenorum Baptista and Mathis 
6. Face of d and @ angulate in profile, sloped 
anteroventrally from base of antenna to vi- 
brissal angle; facial carina lacking in 6; 6 
frons entirely depressed below insertion of 
fronto-orbital setae; eye densely microsetu- 
lose (Colombia) 
C. colombianus Baptista and Mathis 
— Face of ¢ shallowly and vertically arched, 
never angulate; facial carina present in d; d 
frons depressed only medially; eye microse- 
tulae sparse, difficult to discern 
7. Wing cell R,,; usually completely infuscate, 
dividing subapical white spot (pale specimens 
with only traces of brown on the veins around 
cell R,,;); mid- and hindfemora mostly yel- 
LOWE SE in be BBWS eee 8 
— Wing cell R,,; with a conspicuous, undivided 
subapical white spot; mid- and hindfemora 


mostly dark (a5 Weds ctl a a ee 11 
8. Basal rays of arista not bifurcate; middle por- 

tion of hindtibia darkened; postpronotum 

shiny to subshiny, microtomentose ....... 9 


VOLUME 102, NUMBER 3 


— Basal 3—4 rays of arista bifurcate; hindtibia 
completely yellow; postpronotum shiny, lack- 
ing microtomentum (Costa Rica, Mexico) ... 

C. americus Baptista and Mathis 

9. Forefemur brownish dorsally; vertex with a 
large, shiny spot; surstylus as in Fig. 1 (Guy- 
ATLA) Bo opto Scorch oreqeeee Sale C. funkae, new species 

— Forefemur mostly yellow dorsally; vertex en- 
tirely subshiny microtomentose or with a tiny 
shiny spot adjacent to posterior ocellus; sur- 
stylus otherwise 

10. Hindfemur infuscate on apical 1/6; surstyli 
very elongate, apical portion concealed below 
6th—7th sternite; right surstylus boot-shaped; 
produced portion of male face % width of Ist 
haccllomerce ei sic cee oe Oss C. sp. 1 

— Hindfemur infuscate on apical %; surstyli not 
elongate, apex of right surstylus only partially 
concealed below 6th—7th sternite; right sur- 
stylus as in Fig. 7; male face only slightly 
produced, produced portion of face %4 width 
of 1st flagellomere Gsspx2 

11. Right surstylus with apical margin deeply 
concave in middle, having a hook-shaped ap- 
pearance; aedeagal apodeme length equal to 
length of combined 6th and pregenital tergites 
(Baptista and Mathis 1996:246, Fig. 1) (Bra- 
Al!) uments cae C. sabroskyi Baptista and Mathis 

— Right surstylus with apical margin convolut- 
ed, irregular (Baptista and Mathis 1994:24, 
Fig. 58); aedeagal apodeme very large, length 
equal to combined length of 5th, 6th, and pre- 
genital tergites (Brazil, Costa Rica, Peru) 

C. neotropicus Hennig 


Cyamops funkae Baptista and Mathis, 
new species 
(Figs. 1-4, 32) 


Baptista and Mathis 1996: 247 [misidenti- 
fied as C. americus Baptista and Mathis]. 


Description.—Adult d length 2.5—2.6 
mm; wing length 2.1—2.4 mm; wing width 
0.8 mm. Adult 2 length 3.0 mm; wing 
length 3.5 mm; wing width 1.0 mm. 

Head: Ocellar tubercle polished; vertex 
with a large shiny spot; remaining frons 
subshiny, microtomentose, velvet at deepest 
portion; antenna mostly yellow, infuscate at 
dorsal margin; facial region intensely whit- 
ish microtomentose, brownish dorsally; pal- 
pus and labellum mostly white; face in pro- 
file sloped ventrally from base of antenna 
to vibrissal angle, then slightly receded to 
oral margin, produced portion of face same 


483 


width as medial portion of Ist flagellomere; 
eye sparsely microsetulose; arista bearing 
about 10-12 dorsal, 3 ventral rays, none 
clearly bifurcate. Chaetotaxy: Inner fronto- 
orbital setae slightly divergent, almost same 
length as outer vertical seta; pseudovibris- 
sae divergent, curved dorsally; peristomal 
setae 7—10, intercalated by a few setulae. 

Thorax: Postpronotum sparsely microto- 
mentose; halter mostly white, brown at 
base; scutellum trapezoidal, orientation of 
scutellum slightly more elevated than scu- 
tum, disc flattened. Chaetotaxy: Dorsocen- 
tral setae 0+ 1, same length as basal scutel- 
lar seta; some setulae between dorsocentral 
and acrostichal rows; setulae of mesonotum 
short and numerous; scutellar setae 2, basal 
seta about % length of apical seta; anepis- 
ternum with 2-3 posterior setae. Legs: 
Mostly yellow; forecoxa pale yellow, whit- 
ish microtomentose; midcoxa strongly mi- 
crotomentose; forefemur mostly brown dor- 
sally; apical %4 of hindfemur, apical and 
sometimes subapical tarsomere of each leg 
brown; hindtibia darkened on central por- 
tion; spinelike setulae 5—7, weakly differ- 
entiated and difficult to discern. Wing (Fig. 
32): Partially hyaline, mostly with dark pat- 
tern; cell R,,; usually entirely infuscate, di- 
viding subapical white spot; Ist costal ratio: 
1.2-1.37 (wing not slide-mounted); 2nd 
costal ratio 2.97-3.33 (wing not slide- 
mounted); wing ratio 2.7—3.5. 

Abdomen (Figs. 1—3): All segments sub- 
shiny microtomentose; 7th tergite as wide 
as 6th; 6th—7th sternite as in Fig. 2, setulose 
along postarior portion. Terminalia (Figs. 1, 
3): Left surstylus long, slightly inclinate at 
apical %4, posteroapical portion apparently 
excavated, although without a true concav- 
ity; right surstylus shorter and broader than 
left; ejaculatory apodeme (Fig. 3) small, al- 
most equal to length of 5th tergite (a little 
longer than combined 6th and pregenital 
tergites), in lateral view hatchet shaped, ex- 
tended process parallel sided, in dorsal view 
somewhat triangular in shape. 

Female: Head: First flagellomere with 
dorsal half and apex brown; mesofacialia 


484 


Figs. 1—4. 
4—7, ventral view. 3, Internal male terminalia, lateral view. 4, Female abdomen, segments 6—9, ventral view. 


Cyamops funkae. 


Abbreviations: aed. 


epandrium; gon. = gonite; st. = sternite. 


dark brown, shiny medially; gena light 
brown to brown in ground color; palpus 
brown; produced portion of face longer 
than Ist flagellomere medially; arista bear- 
ing 14 dorsal rays. 

Thorax: Dorsocentral setulae typically 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


6th-7th st. 


Hi! Mtg Mi, 


WAAL), 
Say \f 44 
As My 
NO 
> 


1, Epandrium, cerci, and surstyli, ventral view. 2, Male abdominal segments 


= aedeagus; aed. apod. = aedeagal apodeme; ej. apod. = ejaculatory apodeme; ep. = 


more developed than in males. Legs with 
base of forecoxa sometimes mostly black. 
Abdomen (Fig. 4): Width of 7th tergite 
about % that of 6th tergite; 8th sternite ap- 
parently fused to 7th sternite, as in Fig. 4, 
not completely conspicuous, represented by 


VOLUME 102, NUMBER 3 


a pair of lateral setulose lobes and a central 
membranous lobe; 4 spermathecae, round- 
ed, 1 pair slightly smaller than others. 

Type material——The holotype ¢ is la- 
beled “GUYANA. CEIBA (ca.40 km S 
Georgetown) 06°29.9'N, 58°13.1’W{[,] 21 
April 1995[,] Wayne N. Mathis/USNM 
ENT 00134287 [plastic bar code label]/HO- 
LOTYPE 6 Cyamops funkae Baptista & 
W.N.Mathis USNM [red; species name, 
gender symbol, and “Baptista &” hand- 
written]. The holotype is double mounted 
(minuten in a block of plastic), is in excel- 
lent condition, and is deposited in the 
USNM. Paratypes are as follows: Same la- 
bel data as the holotype (2 6; USNM; 1 ¢ 
with abdomen dissected, structures in an at- 
tached microvial); same label data as the 
holotype but with the following changes in 
dates: 13 Apr 1994 (1 2 USNM; abdomen 
dissected, structures in an attached micro- 
vial); 28 Aug 1997 (3 36; 1 2; USNM). 

Distribution.—Cyamops funkae is known 
only from the type locality, CEIBA 
(06°29.9'N, 58°13.1'W), in Guyana. 

Etymology.—The species epithet, fun- 
kae, is a patronym to honor, recognize, and 
express appreciation to Dr. Vicki A. Funk, 
Director of the Biodiversity of the Guianas 
Program, who supported and encouraged 
our field work in Guyana. 

Remarks.—In our last paper on Cyamops 
(Baptista and Mathis 1996), we misidenti- 
fied C. funkae as C. americus and reported 
Guyana as a new locality for the latter spe- 
cies. The two species are very similar, and 
care must be taken to distinguish between 
them. Although the dorsal aristal branches 
of C. funkae are not bifurcate, which seems 
to be a consistent character, we relied on 
the structures of the male terminalia to dis- 
tinguish this species. 

The female of C. funkae can easily be 
distinguished from that of C. americus by 
the shape of the 8th sternite, which is ap- 
parently reduced to a pair of lateral setose 
lobes and a well-defined median lobe. Fe- 
males of C. americus have lateral plates 
without setulae and a membranous median 


485 


lobe that is less conspicuous. The 7th ter- 
gite of the female abdomen is very narrow 
in C. americus, approximately “4 the width 
of the preceding tergite, and the spermathe- 
cae are elongate, not rounded as in this spe- 
cies. The shape of the 6th-7th sternite and 
of the surstyli is unique to this species. 


Cyamops sp. 1 
(Figs. 5—6) 

A specimen from Trinidad, representing 
an undescribed species, can be distin- 
guished from other Neotropical species of 
Cyamops by the characters given in the key. 
Although we are not naming this species 
here because the only available specimen is 
in poor condition, we are providing an il- 
lustration of the male terminalia (Figs. 5— 
6). This species is similar to C. sp. 2 (see 
below) but can be distinguished from it by 
the characters given in the key. The shape 
of the right surstylus is boot-shaped and 
that of the 6th—7th sternite is quite distinc- 
tive and apparently unique among species 
of the genus. 

Material examined.—The specimen is la- 
beled ‘Trinidad 20 Jun [date handwritten] 
WI/Aug. Busck Collector” (1 6; USNM). 


Cyamops sp. 2 
(Fig. 7) 

This unique male from Panama appar- 
ently represents a new species. We defer 
formal description for the same reasons just 
noted (see Cyamops sp. 1). An illustration 
of the male terminalia is provided, however 
(Fig. 7). This species is most similar to C. 
sp. 1, described above, but can be easily 
distinguished from that species by the char- 
acters given in the key. The shape of the 
right surstylus and of the 6th-7th sternite is 
unique for this species. 

Material examined.—The specimen is la- 
beled “‘LaJollaPan[ama] 29 ITV 1952 [29 
Apr 1952]/FSBlanton Collector” (1 6; 
USNM). 


AFROTROPICAL REGION 


Until now, no species had been described 
from this region, although Sabrosky (1980) 


486 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


i 


Figs. 5-7. Cyamops sp. 1 (Trinidad). 5, Segments 5—7, epandrium, cerci, and surstyli, ventral view. 6, 


Ejaculatory apodeme, lateral view. Cyamops sp. 2 (Panama). 7, Segments 5—7, epandrium, cerci, and surstyli, 
ventral view. 


°° 


listed a “sp.” from Nigeria and Khoo KEry TO AFROTROPICAL SPECIES OF CYAMOPS 
(1985) mentioned an ‘undescribed spe- 


Sane, ; 1. Scutellar setae 1 pair; dorsocentral setae 0 + 
cies”? from Africa. 


2; hindfemur mostly yellow, infuscate apically; 


Both of the Afrotropical species de- wing mostly brown with 2 conspicuous white 
scribed below are characterized by the fol- spots along anterior margin and 2 white spots 
lowing combination of characters: Face of along posterior margin; right surstylus sities 


: : : angular, not claw shaped in appearance (Ni- 
male constricted medially; crossvein bm-cu E Seam in SY 
SELMA)! yee ep ore C. nigeriensis, new species 


DResent; separating cell bm from cell dc; _ scutellar setae 2 pairs; dorsocentral seta 0 + 
anepisternum not setulose posteriorly. 1; hindfemur brown on apical %; wing hyaline; 


VOLUME 102, NUMBER 3 


right surstylus with apical margin deeply con- 
cave in middle, claw shaped in appearance 
(Madagascar) ....... C. freidbergi, new species 


Cyamops nigeriensis Baptista and 
Mathis, new species 
(Figs. 8—10, 33) 


Description.—Adult d length 1.6—2.0 
mm; wing length 1.6—1.7; wing width 0.5— 
0.6 mm. Adult 2 length 1.8—2.0 mm; wing 
length 1.9 mm; wing width 0.6—0.7 mm. 

Head: Vertex shiny, ocellar tubercle and 
surrounding area almost polished; de- 
pressed region of frons deep black; antenna 
yellow, some specimens with dorsal margin 
slightly infuscate; facial region mostly yel- 
low, sparsely microtomentose medially; la- 
bellum and palpus pale yellow; face shal- 
lowly and vertically arched, not angulate, 
slightly produced, produced portion about 
% width of Ist flagellomere. Chaetotaxy: 
Inner fronto-orbital setae nearly parallel to 
slightly divergent, %—%4 length of outer ver- 
tical seta; arista bearing 9—10 dorsal, 3 ven- 
tral rays, 6—7 basal rays clearly bifurcate; 
pseudovibrissae slightly converging, orient- 
ed dorsally; peristomal setae 3—4, some se- 
tulae intercalated between them. 

Thorax: Halter mostly white; scutellum 
more or less trapezoidal, apex rounded, ori- 
entation of scutellum slightly more elevated 
than scutum, disk slightly convex; post- 
pronotum narrow and shiny; anepisternum 
shiny; anepimeron polished. Chaetotaxy: 
Dorsocentral setae 0 + 2, posterior seta 
subequal in length to scutellar seta, anterior 
dorsocentral seta slightly shorter and thin- 
ner; mesonotal setulae very thin and sparse; 
scutellar seta 1. Legs mostly yellow; mid- 
and hindfemora infuscate, becoming darker 
brown apically; apical tarsomere of each leg 
brown; spinelike setulae not differentiated. 
Wing (Fig. 33): Mostly with dark pattern; 
cell R, mostly hyaline; cell R,,; with a bas- 
al and subapical oblique spot; cell R,,; 
completely brown; cell M with a mediob- 
lique spot; posterior margin of wing with 2 
well-defined, hyaline spots (some speci- 
mens with an additional hyaline spot at anal 


487 


angle); vein R,,; strongly sinuous about 
midlength; Ist costal ratio: 1.3—2.22 (slide- 
mounted wing 1.42); 2nd costal ratio 3.84— 
5.0 (slide-mounted wing 5.0); wing ratio 
2.42—2.57 (slide-mounted wing 2.5). 

Abdomen (Figs. 8—9): Abdominal seg- 
ments subshiny; 6th—7th sternite as in Fig. 
8. Male terminalia (Figs. 8-9): Right sur- 
stylus large, slightly longer than wide, al- 
most same length as left; left surstylus nar- 
row, gradually tapered to point, internal 
margin slightly sinuous, approximately 
boot-shaped; ejaculatory apodeme subequal 
in length to 5th tergite, in lateral view 
hatchet shaped, extended process parallel 
sided, in dorsal view somewhat tear drop in 
shape. 

Female: Head: Mesofacialia and gena 
dark brown, strongly microtomentose, me- 
dial portion of face and clypeus shiny; pal- 
pus brown; antenna dorsally infuscate; face 
in profile sloped anteroventrally from base 
of antenna to vibrissal angle, thereafter 
slightly receded to oral margin. 

Thorax: Setae and setulae of mesonotum 
typically better developed and longer than 
in male. Legs with foretarsi brown; apical 
%—Y of midfemur brown; apical 4—% of 
hindfemur brown. 

Abdomen (Fig. 10): 7th tergite and ster- 
nite forming a complete ring as in Ameri- 
can species, widened ventrally with poste- 
rior margin slightly pointed, very thin dor- 
sally, not well delimited and concealed be- 
neath 6th tergite; 2 spermathecae, apical 
portion of spermathecal duct apparently un- 
sclerotized. 

Type material—The holotype ¢ is la- 
beled “‘Nigeria Badeggi Rice Res. Sta., NW 
state12-22-68 [22 Dec 1968]/J.T.Medler 
Collector/HOLOTYPE 3 Cyamops niger- 
iensis Baptista & W.N.Mathis [red; species 
name, gender symbol, and “Baptista &” 
handwritten].’’ The holotype is double 
mounted (glued to a point), is in relatively 
good condition (setae of face and thorax 
half broken, following tarsi missing; fore 
right, mid right, hind left), and is deposited 
in the USNM. Paratypes are as follows: 


488 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 8-10. Cyamops nigeriensis. 8, Segments 4—7, epandrium, cerci, and surstyli, ventral view. 9, Epan- 
drium, internal male terminalia, lateral view. 10, Female abdomen, segments 5—9, dorsal view. 


Same locality label as the holotype (4 6, 8 
2; none in good condition; 1 6, 1 2 with 
abdomen dissected, structures in an _ at- 
tached microvial; USNM). 

Other material examined.—Nigeria. Za- 
ria, Samaru, 23 Jun 1966, J. C. Deeming (2 
2; USNM). 


Distribution.—Cyamops nigeriensis is 
known only from Nigeria. 

Etymology.—The species epithet, niger- 
iensis, refers to the country where the type 
locality is located. 

Remarks.—A character that distinguishes 
this species from C. freidbergi, the presence 


VOLUME 102, NUMBER 3 


of a single pair of scutellar setae, also oc- 
curs in C. imitatus Sturtevant (North Amer- 
ica). The narrowed postpronotum is appar- 
ently unique to C. nigeriensis. 


Cyamops freidbergi Baptista and Mathis, 
new species 
(Figs. 11-13) 


Description.—Holotype ¢ length 1.9 
mm; wing length 1.74; wing width 3.2 mm. 

Head: Vertex almost completely pol- 
ished; ocellar tubercle shiny; depressed re- 
gion of frons deep black; antenna yellow, 
pedicel infuscate dorsally; facial region yel- 
low, sparsely microtomentose medially; 
palpus and labellum pale yellow; face shal- 
lowly and vertically arched, not angulate, 
only slightly produced, produced portion al- 
most width of Ist flagellomere. Chaetotaxy: 
Inner fronto-orbital setae nearly parallel to 
slightly divergent, about half length of outer 
vertical seta; arista bearing 10 dorsal, 3 ven- 
tral rays, 6 dorsobasal rays clearly bifur- 
cate; pseudovibrissae slightly convergent, 
oriented dorsally; peristomal setae 9, with 
longest seta about half length of pseudovi- 
brissal seta. 

Thorax: Halter pale yellow; scutellum 
trapezoidal, orientation of scutellum slight- 
ly more elevated than scutum, disk flat; 
postpronotum shiny, slightly microtomen- 
tose. Chaetotaxy: Dorsocentral setae 0+1; 
mesonotal setulae very thin and sparse, 
mostly restricted to dorsocentral and acros- 
tichal rows; scutellar setae 2, basal pair 
about % length of apical seta. Legs mostly 
yellow; hindfemur brown on apical %; api- 
cal and subapical tarsomere of each leg 
brown; spinelike setulae weakly differenti- 
ated, brown, 5. Wing: Hyaline, central por- 
tion very slightly infuscate with brown; Ist 
costal ratio 1.2 (slide-mounted wing); 2nd 
costal ratio 3.75 (slide-mounted wing); 
wing ratio 4.16 (slide-mounted wing). 

Abdomen (Figs. 11—13): 6th tergite as 
large as 7th tergite dorsally. Male terminalia 
(Figs. 11, 13): Right surstylus short, almost 
as long as wide, about a half length of left 
surstylus, apical margin deeply concave in 


489 


middle, hook-shaped in appearance; left 
surstylus narrow, gradually tapered to point, 
margins slightly sinuous, basally slightly 
expanded to left; ejaculatory apodeme in 
lateral view hatchet shaped, extended pro- 
cess parallel sided, in dorsal view some- 
what tear drop in shape. 

Female: Unknown. 

Type material——The holotype ¢ is labeled 
“MADAGASCAR NWI,] NosyBe, Forest 
SE[,] Lakobe Res[,] 5.IV.1991 [5 Apr 1991] 
A. FREIDBERG & FINI KAPLAN/PERIS- 
CELIDIDAE Cyamops sp. D.K. McAlpine 
det. 1991 [handwritten except for ‘“‘D. 
McAlpine Det.” ]/Dissected by A.Baptista 97 
[black border]J/HOLOTYPE 3d Cyamops 
freidbergi Baptista & W.N.Mathis [red; spe- 
cies name, gender symbol, and “Baptista &”’ 
handwritten].’’ The holotype is double 
mounted (minuten in a block of plastic), is in 
relatively good condition (abdomen removed, 
dissected, structures in an attached microvi- 
al), and is deposited in the USNM. 

Distribution.—This species is known 
thus far only from the type locality on Mad- 
agascar. 

Etymology.—tThe species epithet, freid- 
bergi, is a genitive patronym to honor and 
recognize Dr. Amnon Freidberg, who col- 
lected this species and many other interest- 
ing acalyptrate Diptera on Madagascar. 

Remarks.—The presence of a single dor- 
socentral seta, a character that distinguished 
this species from C. nigeriensis, also occurs 
in New World species of Cyamops. The 
shape of the male surstylus (Fig. 11) distin- 
guishes this species from other congeners. 


AUSTRALASIAN/OCEANIAN REGION 


The Australasian/Oceanian fauna demon- 
strates variation in characters that are con- 
stant in American and Afrotropical species, 
i.e., sexual dimorphism in the shape of the 
face (absent in C. micronesicus and an un- 
described species from Ponape) and pres- 
ence of crossvein bm-cu. All species, how- 
ever, have two dorsocentral setae, and the 
anepisternum lacks setae along the posterior 
Margin. 


490 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


KEY TO AUSTRALIAN/OCEANIAN SPECIES OF 


. Wing hyaline (Australia) . 


CYAMOPS 


‘wBasdlatistal rays not bilurcate =. s.cy. meee 2 
Basalvaristal rays biturcate =~. 2-3. -s- eaemer =) 

. Mesofacialia bearing a ridge (male) or a wide 
elevated portion (female); face of male nar- 


row, constricted medially; 1st costal ratio 2.3 
C. fiji, new species 


OL sreater (Eiji)... - amen 
Mesofacialia flat throughout; face of male 


wide, not constricted medially; 1st costal ratio 


PORN S ee: wy cho tteeen. ie ere et eh Seat 3 
. Pseudovibrissa aligned with other peristomal 

setae; basal scutellar seta about % length of 

apicalsSetalg sa. tc < Ree he ee eee ee 4 


Pseudovibrissa placed externally to the row of 
peristomal setae; basal scutellar seta about %4 

or more length of apical seta (Micronesia) 
“Ponape” species complex 


. Mesofacial plate without setae; tibia and tar- 


sus of foreleg mostly brown to dark-brown; 
ocellar tubercle shiny; vertex shiny (Yap) 
Lea es. a eae C. micronesicus, new species 
Mesofacial plate setose between upper peri- 
stomal setae; tibia and tarsus of foreleg most- 
ly yellow; ocellar tubercle dull microtomen- 


tose; vertex dull microtomentose (Ponape) . . . 


Sah cae Meche Ag igh te Uh eS ecaend Po MPEIE dln vada (Cs SF 3 
istreostalsrationlO=leGieee see, eee ee 6 
listacostaliratio) 225.0 mMorem eee ener 10 


Comb on ventral margin of midcoxa present 


(Australia) C. pectinatus Khoo 


. Peristomal setae on mesofacial plate (Austra- 


arene sac. 3 ORS te C. claudiensis Khoo 
Peristomal setae on genal suture......... 8 
. Basal scutellar seta about %4 or more length of 
apical seta; tibia and tarsus of foreleg mostly 
yellow to yellowish brown............. 9 


Basal scutellar seta at most % length of apical 
seta; tibia and tarsus of foreleg mostly brown 
to dark-brown (New Guinea) 


C. papuensis, new species 
. C. truncatus Khoo 
Wing with a conspicuous brown pattern (Aus- 

tralia) C. dayi Khoo 
wh, AEE: 11 
Sth sternite of male abdomen divided medi- 

ally (American Samoa) 


C. samoensis, new species 


- Legs mostly yellowish; midfemur yellow; 


wing hyaline (Australia) C. australicus Hennig 
Legs mostly yellowish brown to black; mid- 
femur brown apically; wing with a conspic- 


uous brown pattern (Australia) .. C. delta Khoo 


Cyamops micronesicus Baptista and 
Mathis, new species 
(Figs. 14-16) 


Description.—Adult ¢ length 2.0 mm; 
wing length 1.8 mm; wing width 0.6 mm. 
Adult ? length 1.8—2.07 mm; wing length 
1.97—2 mm; wing width 0.67—0.72 mm. 

Head: Vertex and ocellar tubercle bright 
shiny, depressed region of frons subshiny; 
antenna yellow; facial region yellow in 
ground color, gray microtomentose; clypeus 
and palpus yellow; face shallowly and ver- 
tically arched, not angulate, nor constricted 
medially. Chaetotaxy: Inner fronto-orbital 
setae nearly parallel to slightly divergent, 
slightly smaller than outer vertical seta; 
arista bearing 9—10 dorsal, 3 ventral rays, 
none clearly bifurcate; pseudovibrissae 
slightly divergent, pointing dorsally; peri- 
stomal setae 6—7. 

Thorax: Halter mostly pale yellow; scu- 
tellum trapezoidal, orientation of scutellum 
very slightly more elevated than scutum, 
disk slightly convex; postpronotum and up- 
per margin of notopleuron shiny. Chaeto- 
taxy: Dorsocentral setae 0+2, posterior seta 
subequal in length to apical scutellar seta 
but thinner, anterior dorsocentral seta slight- 
ly shorter and thinner; mesonotal setulae 
very thin and sparse; scutellar setae 2, basal 


seta about % of apical seta. Legs mostly yel- 


low (foretibia and foretarsi of holotype 
missing); apical %4—% of hindfemur, apical 
and subapical tarsomere of each leg brown; 
spinelike setulae not differentiated. Wing: 
Partially hyaline; cell R,,; mostly brown, 
subdividing subapical spot; cell M and 
somewhat cell CuA, mostly lightly infus- 
cate to hyaline; vein R,,,; only slightly sin- 
uous at midlength; cells bm and dm conflu- 
ent; Ist costal ratio 1.15—1.4 (slide-mounted 
wing); 2nd costal ratio 2.2—3.8 (slide- 
mounted wing); wing ratio 2.88—2.96 
(slide-mounted wing). 

Abdomen (Figs. 14—16): 6th tergite sub- 
equal in width to 7th tergite. Male termin- 
alia (Figs. 14-15): Left surstylus narrow, 
gradually tapered to a point, about 1.5 times 


VOLUME 102, NUMBER 3 491 


aed. 
Figs. 11-13. Cyamops freidbergi. 11, Epandrium, cerci, and surstyli, ventral view. 12, Male abdominal 
segments 5—7, ventral view. 13, Internal male terminalia, lateral view. Abbreviations: aed. = aedeagus; aed. 


apod. = aedeagal apodeme; gon. = gonite; st. = sternite. 


longer than right; right surstylus slightly view as long as 6th tergite, with extended 
longer than wide, anterior margin slightly process greatly expanded toward apex, fan- 
receded, forming a nipplelike projection on _ like (Fig. 16). 

inner corner; ejaculatory apodeme in dorsal Female: Head: Mesofacialia and gena 


492 


16 


Figs. 14-16. 


COR ses eS 
urs 
am A) 


lS 


Cyamops micronesicus. 14, Epandrium, cerci, and surstyli, ventral view. 15, Male abdominal 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


oN ‘ (HY 


segments 4—7, ventral view. 16, Ejaculatory apodeme, lateral view. 


brown in ground color, strongly invested 
with grayish microtomentum; inner margin 
of pedicel dorsally infuscate. 

Thorax: Setae and setulae of mesonotum 
typically better developed and longer than 
in male; postpronotum almost lacking mi- 
crotomentum; tibia and tarsus of foreleg 
brown to dark brown. Wing: pattern gen- 
erally darker than in male. 

Abdomen: 7th tergite and sternite form- 
ing a complete ring, wide ventrally, dorsally 
about half that of 6th tergite; 2 subequal 
spermathecae. 


Type material—The holotype ¢ is la- 
beled ‘“‘YaptownYap VII-13 -46 [13 Jul 
1946]/HKTownes No 1090 [number hand- 
writtenJ/HOLOTYPE Cyamops micronesi- 
cus Baptista & W.N.Mathis USNM [red; 
species name, gender symbol, and “*Baptis- 
ta &’’ handwritten].”’ The holotype is dou- 
ble mounted (glued to a point on ventral, 
right side), is in fair condition (foretibiae 
and tarsi missing, left wing detached and 
slide-mounted; abdomen removed, dissect- 
ed, structures in an attached microvial), and 
is deposited in the USNM. Paratypes are as 


VOLUME 102, NUMBER 3 


follows: Same locality label as the holotype 
but with different dates: 12, 14 Jul 1946 (3 
2; USNM). 

Distribution.—This species is known 
only from the type locality in Micronesia. 

Etymology.—tThe species epithet, micro- 
nesicus, refers to the island group in the 
South Pacific where this species was col- 
lected. 

Remarks.—This species is similar to the 
other Micronesian species, the underscribed 
species from the ‘“‘Ponape”’ complex, and to 
C. sp. 3. It is easily distinguished from 
these species by having the pseudovibrissae 
aligned with the other peristomal setae and 
in lacking setae on the mesofacialia. 


Cyamops fiji Baptista and Mathis, 
new species 
(Figs. 17-19, 34) 


Description.—Adult ¢ length 1.54—1.7 
mm; wing length 1.6—-1.68 mm; wing 
width 0.5—0.6 mm. Adult @ length 1.8— 
1.86 mm; wing length 1.9 mm; wing width 
0.7—0.74 mm. 

Head: Vertex and ocellar tubercle micro- 
tomentose, except for a small shiny dot ad- 
jacent to posterior ocellus; depressed region 
of frons subshiny; antenna yellow; facial re- 
gion yellow in ground color, whitish micro- 
tomentose; clypeus and palpus yellow. 
Chaetotaxy: Inner fronto-orbital setae 
slightly divergent, slightly smaller than out- 
er vertical seta; arista bearing 7 dorsal, 3 
ventral rays, none clearly bifurcate; pseu- 
dovibrissae oriented dorsally; peristomal 
setae 5-7. 

Thorax: Halter mostly dark brown; post- 
pronotum microtomentose, subshiny; scu- 
tellum trapezoidal, orientation of scutellum 
very slightly more elevated than scutum, 
disk slightly convex. Chaetotaxy: Dorso- 
central setae 0+ 1 (some specimens with an 
elongate but thin setula anterior of dorso- 
central seta); mesonotal setulae well devel- 
oped; scutellar setae 2. Legs mostly yellow; 
forefemur a little infuscate dorsally, mostly 
on apical portion; apical 4—% of hindfemur 
brown; apical and sometimes subapical tar- 


493 


somere of each leg brown; spinelike setulae 
not differentiated. Wing (Fig. 34): Hyaline 
to faintly brown with slight traces of brown 
around veins and in cell R,; vein R,,, very 
slightly sinuous at midlength; cells bm and 
dm separated; Ist costal ratio 2.35—2.8 
(slide-mounted wing); 2nd costal ratio 2.1— 
2.5 (slide-mounted wing); wing ratio 2.4— 
3.2 (slide-mounted wing). 

Abdomen (Figs. 17-18): 6th sternite ap- 
parently not completely fused with 7th ster- 
nite. Male terminalia (Fig. 17): Right sur- 
stylus large, slightly longer than wide, ex- 
panded and convolted apically; left sursty- 
lus narrow, gradually tapered to rounded 
apex, slightly bent inward, a little longer 
than right surstylus; aedeagal apodeme with 
a ventral projection joining hypandrium 
posteriorly; ejaculatory apodeme small, 
slightly longer than 7th tergite, in lateral 
view hatchet shaped, extended process par- 
allel sided and slightly expanded at apical 
portion, in dorsal view somewhat subrect- 
angular in shape with margins slightly con- 
verging toward apex. 

Female: Head: Mesofacialia and gena 
brown, strongly microtomentose, silver ex- 
cept on lower central portion of mesofaci- 
alia, where microtomentum is thin; inner 
margin of pedicel dorsally infuscate. 

Thorax: Setae and setulae of mesonotum 
typically better developed and longer than 
in male. Legs with midfemur a little infus- 
cate dorsoapically; hindfemur usually infus- 
cate on apical half or slightly more. Wing: 
Cells R, and in some specimens R,,,; infus- 
cate, brown. 

Abdomen (Figs. 19): 7th tergite and ster- 
nite forming a complete ring, wide ventral- 
ly, dorsally about %4 of 6th tergite; 3 sube- 
qual spermathecae (Fig. 19); sclerotized 
portion of spermathecai duct about same 
length as spermatheca. 

Type material—The holotype ¢ is la- 
beled “FIJI: Viti Levu I: Lami, 0-200 m, 
Ill.1981/N.L.H. Krauss, Coll. BISHOP Mu- 
seum Acc. #1981.131/HOLOTYPE ¢ Cy- 
amops fiji Baptista & W.N.Mathis [red; spe- 
cies name, gender symbol, and “Baptista 


494 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 17-19. 
4—7, ventral view. 19, Spermathecae, ventral view. 


&” handwritten].”” The holotype is double 
mounted (glued to a point on the right side), 
is in relatively good condition (mesonotal 
setae broken), and is deposited in the 
BPBM. Paratypes are as follows: Same la- 
bel data as the holotype (17 6, 18 @; 
BPBM, USNM). 

Etymology.—The species epithet, fiji, is 


Cyamops fiji. 17, Epandrium, cerci, and surstyli, ventral view. 18, Male abdominal segments 


the name of the country where the type lo- 
cality is located and is a noun in apposition. 

Remarks.—This species has the 6th and 
7th sternites incompletely fused, a character 
shared with C. samoensis. Both species, 
however, can be easily distinguished by the 
characters in the key. The shape of the right 
surstylus is unique to C. fiji. 


VOLUME 102, NUMBER 3 


Cyamops papuensis Baptista and 
Mathis, new species 
(Fig. 20) 


Description.—Adult ¢ length 1.7—1.8 

mm; wing length 1.6 mm; wing width 
1.6—2 mm. Adult 2 length 1.8-2.0 mm; 
wing length 1.8—2.0 mm; wing width 2.5 
mm. 
Head: Ocellar tubercle polished; shiny 
spot on vertex large and distinct, extended 
from ocellus % distance to eye margin; de- 
pressed region of frons deep, velvet; pedicel 
on upper half, 1st flagellomere yellow, in- 
fuscate dorsally; facial region yellow in 
ground color, strongly microtomentose; la- 
bellum and palpus pale yellow; face pro- 
duced and slightly angulate. Chaetotaxy: 
Inner fronto-orbital setae slightly divergent, 
slightly smaller than outer vertical seta; 
arista bearing 9 dorsal, 3 ventral rays, 6 
basal rays bifurcate; pseudovibrissae ori- 
ented dorsally; peristomal setae 7. 

Thorax: Halter brown; scutellum trape- 
zoidal, orientation of scutellum moderately 
more elevated than scutum, disk a little 
convex; postpronotum polished. Chaeto- 
taxy: Dorsocentral setae 0+2, posterior seta 
slightly longer than anterior seta; mesonotal 
setulae moderately well-developed; scutel- 
lar setae 2, basal seta % length of apical 
seta. Legs mostly yellow; femora brown in- 
fuscate at apical %—% tibiae very slightly 
infuscate; apical and subapical tarsomere of 
each leg brown; spinelike setulae about 6, 
weakly differentiated. Wing: Hyaline, 
slightly fuscous; cells bm and dm separat- 
ed; Ist costal ratio 1.28—1.5 (slide-mounted 
wing); 2nd costal ratio 3.5—4.0 (slide- 
mounted wing); wing ratio 0.8—1.0 (slide- 
mounted wing). 

Abdomen (Fig. 20): 6th tergite about 
same width as dorsal portion of 7th tergite, 
both sclerites almost without setae; 4th and 
5th sternites with well-developed lateral se- 
tae and a row of setae along posterior mar- 
gin, 5th sternite slightly reduced in length, 
posterior margin somewhat receded. Male 
terminalia: right surstylus almost globular, 


495 


posteromedial margin deeply excavated, 
forming a fingerlike projection on left por- 
tion of surstylus, left surstylus at least 4X 
longer than wide. 

Female: Head: Mesofacialia and gena 
brown, strongly microtomentose, silver ex- 
cept on facial carina where microtomentum 
is thin; lst flagellomere infuscate dorsally. 

Thorax: Legs with femora mostly brown. 

Abdomen: 7th tergite and sternite sepa- 
rate; 7th tergite about %4 length of 6th ter- 
gite; 2 subequal, spherical spermathecae; 
sclerotized portion of spermathecal duct 
about V5 length of spermatheca. 

Type material—The holotype ¢ is la- 
beled ““NEW GUINEA: NE Morobe Dis- 
trict Mindik, 1,200—1,600 m, IX.1968 [Sep 
1968]/N.L.H. Krauss Collector BISHOP 
MUSEUM/HOLOTYPE ¢ Cyamops pa- 
puensis Baptista & W.N. Mathis [red; spe- 
cies name, gender symbol, and “Baptista 
&”’ handwritten].’’ The holotype is double 
mounted (glued to a paper point), is in good 
condition (right wing missing), and is de- 
posited in the BPBM. Paratypes are as fol- 
lows: NEW GUINEA. NE. Morobe Dis- 
trict, Mt. Kaindi (N Peak; 2,350 m), 1-14 
Sep 1966 (1 ¢6; USNM). 

Other material examined.—Papua New 
Guinea. NE. Wau, Hospital Creek (1,200 m; 
Malaise trap), 6 Apr 1965, J. Sedlasec (1 
?; BPBM); Amok (165 m), 6 Jan 1960, T. 
C. Maa (1 2; BPBM). SE. Western District. 
Oriomo River (3 6; light trap), 4 Aug 1964, 
H. Clissold (1 6; BPBM). 

Etymology.—The species epithet, pa- 
puensis, refers to Papua New Guinea, the 
country where the type locality is located. 

Remarks.—The reduced 5th sternite, the 
presence of well-developed setae laterally 
on the 4th and Sth sternites, and the shape 
of the 6th-7th sternites and surstylus distin- 
guish this species. 


Cyamops samoensis Baptista and 
Mathis, new species 
(Figs. 21—22) 
Description.—Holotype ¢ length 1.8 
mm; wing length 1.7 mm; wing width 0.6 
mm. 


496 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


23 


Figs. 20-23. Cyamops papuensis. 20, Segments 4—7, epandrium, cerci, and surstyli, ventral view. Cyamops 
sampensis. 21, Epandrium, cerci, and surstyli, ventral view. 22, 4th—7th sternites, ventral view. Cyamops sp. 
(“‘Ponape”’ species complex). 23, Head, anterior view. Abbreviations: st. = sternite. 


Head: Ocellar tubercle microtomentose; vet; pedicel brown, Ist flagellomere yellow; 
a shiny spot on vertex large and distinct, facial region yellow in ground color, strong- 
extended from ocellus % distance to eye ly microtomentose; labellum and palpus 
margin; depressed region of frons deep vel- yellow; face slightly produced, constricted 


VOLUME 102, NUMBER 3 


medially. Chaetotaxy: Inner fronto-orbital 
setae slightly divergent, slightly smaller 
than outer vertical seta; fronto-orbits with 
setulae that run beyond limits of silver-mi- 
crotomentose portion; arista bearing 9 dor- 
sal, 3 ventral rays, 6 basal rays bifurcate; 
pseudovibrissae parallel, oriented dorsally; 
peristomal setae 7. 

Thorax: Halter mostly white, dark brown 
at base; scutellum trapezoidal, orientation 
of scutellum moderately more elevated than 
scutum, disk flat; postpronotum microto- 
mentose. Chaetotaxy: Dorsocentral setae 
0+2, posterior seta slightly longer than an- 
terior seta; mesonotal setulae moderately 
well developed; scutellar setae 2, basal seta 
% of apical seta. Legs mostly yellow; hind- 
femur apically weakly infuscate; apical and 
subapical tarsomere of each leg brown; 
spinelike setulae about 6, weakly differen- 
tiated. Wing: Hyaline to faintly brownish 
infuscate; cells bm and dm separated; Ist 
costal ratio 3.0; 2nd costal ratio 2.4, wing 
ratio 2.8. 

Abdomen (Figs. 21—22): 6th tergite about 
same width as 7th tergite, both tergites al- 
most without setae (in a dry specimen, only 
6 abdominal segments are seen); 5th ster- 
nite asymmetrical, divided in 2 parts; 6th 
sternite apparently incompletely fused with 
7th sternite. Male terminalia: Right sursty- 
lus large, slightly longer than wide, com- 
plicated in shape, convoluted; left surstylus 
narrow, a little enlarged posteriorly, twice 
length of right surstylus; ejaculatory apo- 
deme triangular when viewed dorsally, pos- 
terior process in lateral view parallel-sided, 
not longer than body of apodeme. 

Type material—The holotype ¢ is la- 
beled “‘Amer[ican]. Samoa Tutuila Is. Le- 
one Area/July 27-Aug 5 MR Wheeler 1962 
[date handwrittenJ/ HOLOTYPE 6 Cy- 
amops samoensis Baptista & W.N. Mathis 
USNM [red; species name, gender symbol, 
and “‘Baptista &’’ handwritten].’’ The ho- 
lotype is in good condition (glued to a 
point; abdomen removed and in an attached 
microvial) and is deposited in the USNM. 
Paratype is as follows: SAMOA. Tulia, Na- 


497 


val station, 24 Aug 1940, Sand Zimmerman 
(1 6; BPBM). 

Etymology.—tThe species epithet, sa- 
moensis, refers to the country where the 
type locality is located. 

Remarks.—The shape of the medially di- 
vided, symmetrical 5th sternite and of the 
right surstylus are typical for this species. 
The separated 6th and 7th sternites is a 
character shared with C. fiji. Males of C. 
femoratus also have the 5th sternite divided 
medially. 


Cyamops pectinatus Khoo 


Cyamops pectinatus Khoo 1985: 528— 
530.—Khoo and Sabrosky 1989: 551 
[Australasian/Oceanian catalog]. 


Specimens examined.—AUSTRALIA. 
Tasmania: Stoneyford Creek (N of La- 
goons), 11 Nov 1979, H. B. Williams (1 d; 
USNM). 

Distribution.—North Queensland to the 
southern coast of New South Wales and in- 
land to Narrabri and the Australian Capital 
Territory (Khoo 1985). Tasmania is a new 
state record for this species. 


*‘Ponape”’ species complex 
(Fig. 23) 


Seven specimens from Ponape, all in 
poor condition, share a distinctive charac- 
ter: Pseudovibrissae are placed external to 
the row of peristomal setae (Fig. 23). In ad- 
dition, these species can be easily distin- 
guished from other species from Ponape by 
the absence of mesofacial setae. The male, 
like C. sp. 3, has a wide face. Having per- 
istomal setae that are modified differently 
in the male and the female, along with dif- 
ferences in the coloration of the legs and 
wings, lead us to suspect that the male and 
female might belong to different species. 
Because the wings of available specimens 
are in poor condition, it was impossible to 
determine whether crossvein bm-cu is pre- 
sent or not. Some of the specimens we ex- 
amined came from different localities and 
we are unsure of the number of species rep- 


498 


resented in this series. We hope that study 
of additional and well preserved specimens 
from this region will resolve these prob- 
lems. 

Material examined.—Ponape: Mt. Ta- 
matamnsakis (180 m), 17 Jan 1953 (1 4, 
USNM); E. Caroline Is. Pac. Sci. Bd. J. L. 
Gressitt (2 9, USNM); (N.); SE. Na- 
mpnmal (70 m; light trap), 12 Jan 1953, J. 
L. Gressitt’” (1 2, USNM); Mt Do Len- 
mankap (1,800 ft), 2 Aug 1946 (1 2°; 
USNM); (2,000 ft), 13 Aug 1996 (2 &; 
USNM). 


Cyamops sp. 3 


A unique male specimen, also from Pon- 
ape (see above), is distinguished by having 
long pseudovibrissae (longer than outer ver- 
tical seta) and by having mesofacial setae 
between the uppermost peristomal setae. 
The face is wide, the pseudovibrissae are 
aligned with the peristomal setal row, and 
crossvein bm-cu is present. This is the only 
species of Cyamops known to have meso- 
facial setae. Although this species is distinct 
from the other species occurring on Ponape, 
the specimen is in poor condition and we 
are not formally describing it. 

Material examined.—MICRONESIA. 
Ponape: Mt. Do Lenmankap (1800 ft), 2 
Aug 1946 (1 6; USNM). 


ORIENTAL REGION 


KEY TO ORIENTAL SPECIES OF CYAMOPS 


1. Face of male wide, not constricted medially; 
male and female facialia gray microtomentose, 
bearing a polished triangular region on lower, 
protruded portion (inconspicuous in some 6d 
specimens); forefemur of male with a long bas- 
al ventral seta (Philippines. Negros) ........ 

See Oe ee See POR Ree ae C. femoratus, new species 
— Face of male narrow, constricted medially, fa- 
cialia of male and female without a polished, 
anterior triangle; forefemur without a basal 
ventral seta 

. Basal scutellar seta very thin, about % that of 

apical seta (Thailand) C. kaplanae, new species 

— Basal scutellar seta well developed, about %— 

%4 of apical seta 

3. Hindfemur of male brown at apical 4%; femora 
of female mostly brown; cell R, 


N 


, of wing usu- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ally hyaline on anterior basal portion, other- 

wise brown; right surstylus subrectangular, 

much smaller than left surstylus (Fig. 17) 

(LaOS)) ic. cre kere Se C. banvaneue, new species 
— Hindfemur of male brown at apical 4 femora 

of female mostly yellow; cell R,,; of wing usu- 

ally mostly infuscate dividing subapical white 

spot; right surstyius as in Fig. 26 (Laos) ..... 

Pee rier Cy aoe ale ho oe C. laos, new species 


Cyamops femoratus Baptista and 
Mathis, new species 
(Figs. 24—25, 35) 


Description.—Adult ¢ length 2.3—2.4 
mm; wing length 2.2—2.4 mm; wing width 
0.6-0.7 mm. Adult 2 length 2.4—2.7 mm; 
wing length 2.5—2.6 mm; wing width 0.8— 
0.9 mm. 

Head: Vertex bright shiny, mostly pol- 
ished; ocellar tubercle and depressed region 
of frons subshiny; pedicel brown, Ist fla- 
gellomere yellow; facial region brown in 
ground color, gray microtomentose, anteri- 
orly with a polished triangular region; pal- 
pus and labellum yellow; face shallowly 
and vertically arched below level of pseu- 
dovibrissae, in lateral view as large as wid- 
est portion of 1st flagellomere. Chaetotaxy: 
Inner fronto-orbital setae nearly parallel to 
slightly divergent, slightly smaller than out- 
er vertical seta; arista bearing 12—14 dorsal, 
3-4 ventral rays, none clearly bifurcate; 
pseudovibrissae oriented anteriorly and 
curved laterad; peristomal setae 6—7, 3rd 
and 4th setae better developed and strongly 
inclinate. 

Thorax: Halter mostly brown, base pale 
yellow; scutellum trapezoidal, orientation 
of scutellum very slightly more elevated 
than scutum, disk flat to slightly convex; 
pleural region shiny; postpronotum glossy. 
Chaetotaxy: Dorsocentral setae 0+2, pos- 
terior seta subequal in length to apical scu- 
tellar seta, anterior dorsocentral seta slightly 
shorter and thinner; mesonotal setulae very 
thin and sparse; scutellar setae 2. Legs with 
femora long, almost same length as abdo- 
men; legs mostly yellow; forefemur slightly 
infuscate apically; apical Y10—1/S of hind- 
femur brown; apical and subapical tarso- 


VOLUME 102, NUMBER 3 


499 


Figs. 24-26. Cyamops femoratus. 24, Epandrium, cerci, and surstyli, ventral view. 25, Male S5th—7th sternites, 
ventral view. Cyamops laos. 26, Segments 4—7, epandrium, cerci, and surstyli, ventral view. 


mere of each leg brown; spinelike setulae 
about 4, basalmost seta long, forefemur 
bearing an additional long seta basally. 
Wing (Fig. 35): Mostly hyaline; cells R, 
and R,,, mostly hyaline; posterior margin 
and apex of cell R,,, dark; posterior portion 
of cell M and cell CuA, with hyaline area 
(pale specimens with wing mostly hyaline 
and pattern rather faint); vein R,,, sinuous, 


narrowing cell R,,, at midlength; cells bm 
and dm only partially separated; 1st costal 
ratio 1.3-1.5 (slide-mounted wing); 2nd 
costal ratio 4.3—4.8 (slide-mounted wing); 
wing ratio 2.8—3.6 (slide-mounted wing). 
Abdomen (Figs. 24—25): 5th sternite di- 
vided in middle, slightly asymmetrical; 6th 
tergite almost same width as 7th tergite; 
6th—7th sternites simple, reduced in com- 


500 


parison to other species. Male terminalia 
(Fig. 24): Right surstylus large, slightly lon- 
ger than wide; left surstylus much longer 
than wide, slightly tapered to point; aedea- 
gal apodeme joined to hypandrium poste- 
riorly; length of ejaculatory apodeme in 
dorsal view same as 6th tergite, with ex- 
tended process greatly expanded toward 
apex, fanlike. 

Female: Head: Face silver microtomen- 
tose, lower portion with a polished spot. 

Thorax: Setae and setulae of mesonotum 
typically better developed and longer than 
in male; notopleuron polished dorsad of 
level of setae. Legs with forefemur infus- 
cate brown. 

Abdomen: 7th tergite and sternite form- 
ing a complete ring, wide ventrally, dorsally 
about half width of 6th tergite; 2 subequal 
spermathecae; sclerotized portion of sper- 
mathecal duct subequal to length of sper- 
matheca. 

Type material—The holotype ¢ is la- 
beled ‘‘P[hilippine]. I[slands]., NEGROS 
OR. L. Balinsasayao 6-X-1959 [number 
“6”? handwritten over the top of a printed 
number] /C.M. Yashimoto Collector BISH- 
OP MUSEUM/HOLOTYPE 36 Cyamops 
femoratus Baptista & W.N. Mathis [red; 
species name, gender symbol, and “‘Baptis- 
ta &’’ handwritten].’’ The specimen is dou- 
ble mounted (glued to a paper point), is in 
good condition (left vertical seta missing), 
and is deposited in the BPBM. Paratypes 
are as follows: Same label data as the ho- 
lotype (one 2 with abdomen and wing dis- 
sected and in an attached microvial) but 
with different dates as follows: 6 Oct 1959 
(2 6,4 2; BPBM, USNM); 1-7 Oct 1959 
(3 6,4 2; BPBM). 

Etymology.—tThe species epithet, femor- 
atus, refers to the characteristic seta on the 
forefemur. 

Remarks.—The presence of a basoven- 
tral seta on the male forefemur is an auta- 
pomorphy for this species. The wing pattern 
of C. femoratus resembles that of C. ban- 
vaneue, and a sinuous vein R,,, is shared 
with C. laos and C. nebulosus from North 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


America. The shape of the hypandrium, ae- 
deagal apodeme, and 6th—7th sternite of the 
male abdomen somewhat resembles the 
same structures in C. micronesicus. Like the 
latter species, males of C. samoensis also 
have a medially divided 5th sternite. 


Cyamops laos Baptista and Mathis, 
new species 
(Fig. 26, 36) 


Description.—Adult d length 1.7—1.9 
mm; wing length 1.9—2.0 mm; wing width 
0.64-0.66 mm. Adult 2 length 2.2—2.6 
mm; wing length 2.5—2.8 mm; wing width 
0.86—0.9 mm. 

Head: Ocellar tubercle and vertex almost 
polished; depressed region of frons deep, 
velvet; antenna yellow; face pale yellow, 
strongly microtomentose; labellum and pal- 
pus yellow; face shallowly and vertically 
arched, not angulate, only lower portion 
slightly produced. Chaetotaxy: Inner fronto- 
orbital setae slightly divergent, slightly 
smaller than outer vertical seta; arista bear- 
ing 7-8 dorsal, 3 ventral rays, 6—7 basal 
rays bifurcate; pseudovibrissae slightly di- 
vergent, oriented dorsally; peristomal setae 
6-8. 

Thorax: Halter brown; scutellum trape- 
zoidal, orientation of scutellum moderately 
more elevated than scutum, disk flat; post- 
pronotum polished. Chaetotaxy: Posterior 
dorsocentral seta slightly longer than ante- 
rior seta; mesonotal setulae weakly devel- 
oped; scutellar setae 2, basal seta %4 length 
of apical seta. Legs mostly yellow, only 
hindfemur brown at apical 1; apical and 
subapical tarsomere of each leg brown; 
spinelike setulae about 8, weakly differen- 
tiated. Wing (Fig. 36) mostly slightly infus- 
cate; cell R, mostly hyaline; cell R,,, most- 
ly infuscate with subapical white spot; pos- 
terior portion of cell M and somewhat of 
cell CuA, with hyaline area (wing only 
slightly infuscate around veins in some 
specimens); vein R,,, sinuous, narrowing 
cell R,,; at midlength; Ist costal ratio: 
1.37—1.61 (slide-mounted wing), 1st costal 


VOLUME 102, NUMBER 3 


ratio: 3.25—4.14 (slide-mounted wing), 
wing ratio 2.9—3.1 (slide-mounted wing). 

Abdomen (Fig. 26): 6th tergite about 
same width as 7th, both tergites almost 
without setae; 6th—7th sternite as in Fig. 26. 
Male terminalia (Fig. 26): Right surstylus 
large, longer than wide, gradually tapered 
laterally toward right side; left surstylus 
about 4X longer than wide, much longer 
than right, narrowed toward apex; ejacula- 
tory apodeme short in dorsal view *% as 

Female: Head: Mesofacialia and gena 
dark brown, strongly microtomentose, sil- 
ver except for facial carina which is shiny 
and almost polished on posterior half; ped- 
icel mostly brown; palpus brown to dark 
brown; face angulate, strongly produced, 
produced portion about same width as Ist 
flagellomere. 

Thorax: Wing (Fig. 36): Pattern darker; 
cell R,,; mostly brown except for a hyaline 
median spot. 

Abdomen: 7th tergite and sternite form- 
ing a complete ring, wide ventrally, dorsally 
ranging % of 6th tergite; 2 spermathecae, 
one slightly larger than the other; sclero- 
tized portion of spermathecal duct about 
half length of spermatheca. 

Type material—The holotype ¢ is la- 
beled .““LAOS: Vientiane Prov. Ban Van 
Eue 15.1X.1967 [15 Sep 1967]/Native Col- 
lector BISHOP [Museum]/HOLOTYPE ¢ 
Cyamops laos Baptista & W.N. Mathis [red; 
species name, gender symbol, and “‘Baptis- 
ta &”’ handwritten].’’ The holotype is dou- 
ble mounted (glued to a paper point), is in 
fair condition (facial setae, right Ist flagel- 
lomere, thoracic setae, right hindleg miss- 
ing), and is deposited in the BPBM. Para- 
types are as follows: Same locality label as 
the holotype, but with different dates as fol- 
lows: 15 May 1966 (2 2; BPBM), 8 Aug 
1966 «(12 -BPBM), <15;Aug 1966 Guo; 
BPBM), 15 May 1967 (1 36, 1 2; BPBM, 
USNM), 30 Jun 1967 (2 6, 1 °; BPBM, 
USNM), 15, 30 Aug 1967 (1 2; BPBM), 
15 Sep 1967 (1 6, 1 2; BPBM). One 6 
paratype has been dissected (the structures 
are in an attached microvial). 


501 


Etymology.—The species epithet, laos, 
refers to the country of the type locality and 
iS a noun in apposition. 

Remarks.—This species is similar to C. 
banvaneue but can be easily distinguished 
by the characters cited in the key (also see 
comments under C. banvaneue). 


Cyamops banvaneue Baptista and 
Mathis, new species 
(Figs. 27-30, 37) 


Description.—Adult d length 1.5—2.5 
mm; wing length 1.6 mm; wing width 0.56 
mm. Adult 2 length 1.8—-2.1-mm; wing 
length 1.6—2.1 mm; wing width 0.6—0.7 
mm. 

Head: Ocellar tubercle and vertex slight- 
ly microtomentose to polished, bright 
shiny; depressed region of frons dull; an- 
tenna yellow, pedicel slightly infuscate on 
dorsal margin; face pale yellow, strongly 
microtomentose; labellum and palpus yel- 
low; face shallowly and vertically arched, 
not angulate, only slightly produced, pro- 
duced portion of face about same length as 
width of Ist flagellomere. Chaetotaxy: In- 
ner fronto-orbital setae slightly divergent, 
slightly smaller than outer vertical seta; 
arista bearing 9—10 dorsal, 2 ventral rays, 6 
basal rays bifurcate; pseudovibrissae ori- 
ented dorsally; peristomal setae 7-9. 

Thorax: Halter brown; scutellum trape- 
zoidal, orientation of scutellum slightly 
more elevated than scutum, disk flat to 
slightly convex; postpronotum sparsely mi- 
crotomentose to polished. Chaetotaxy: Pos- 
terior dorsocentral seta slightly longer than 
anterior seta; scutellar setae 2, basal seta 74— 
% length of apical seta. Legs mostly yellow, 
only hindfemur brown at apical half; apical 
and subapical tarsomere of each leg brown; 
forefemur slightly infuscate dorsally; spine- 
like setulae slightly differentiated, 2—3, 
black. Wing (Fig. 37): Mostly infuscate; 
cells R, completely and R,,; mostly hya- 
line; posterior margin and apex of cell R,,; 
dark, posterior portion of cell M and some- 
what cell CuA, with hyaline areas; cells bm 
and dm separated; Ist costal ratio 1.2—1.3 


502 


aed.apod. 


29 


aed. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 27-30. Cyamops banvaneue. 27, Epandrium, cerci, and surstyli, ventral view. 28, Male 6th—7th ster- 
nites, ventral view. 29, Internal male terminalia, lateral view. 30, Spermathecae, ventral view. Abbreviations: 
aed. = aedeagus; aed. apod. = aedeagal apodeme; ej. apod. = ejaculatory apodeme; hy. = hypandrium. 


(slide-mounted wing); 2nd costal ratio 3.7— 
3.8 (slide-mounted wing); wing ratio 2.3— 
2.8 (slide-mounted wing). 

Abdomen (Figs. 27-29): 6th tergite about 
same width as 7th tergite, both tergites al- 
most without setae; 6th—7th sternite as in 
Fig. 28. Male terminalia (Figs. 27, 29): 


Right surstylus large, slightly wider than 
long, subrectangular; left surstylus slightly 
longer than wide, with a fingerlike process 
on apicomesal inner corner about % of 
length of surstylus; aedeagal apodeme 
joined to hypandrium posteriorly; ejacula- 
tory apodeme short in dorsal view at most 


VOLUME 102, NUMBER 3 


% as long as 7th tergite, extended process 
thin, with margins subparallel, in dorsal 
view subrectangular in shape. 

Female: Head: Mesofacialia and gena 
dark brown, strongly microtomentose, sil- 
ver except for facial carina which is shiny 
and almost polished on posterior 74; pedicel 
mostly brown; palpus brown to dark brown; 
femora dark except for yellow basal ring. 

Thorax: Wing: Pattern darker, cell R,,; 
mostly brown except for a hyaline median 
spot. 

Abdomen (Fig. 30): 7th tergite and ster- 
nite forming a complete ring, wide ventral- 
ly, dorsally ranging % of 6th tergite; 2 
spherical subequal spermathecae (Fig. 30); 
length of sclerotized portion of spermathe- 
cal duct about % diameter spermatheca. 

Type material—The holotype ¢ is la- 
beled “LAOS: Vientiane Prov. Ban Van 
Eue 30.VI.1967 [30 Jun 1967] /Native col- 
lector BISHOP [Museum] /HOLOTYPE ¢d 
Cyamops banvaneue Baptista & W.N. 
Mathis [red; species name, gender symbol, 
and “Baptista &”’ handwritten].’’ The ho- 
lotype is double mounted (glued to a paper 
point), is in fair condition (setae of head 
broken, both Ist flagellomeres and many 
thoracic setae missing), and is deposited in 
the BPBM. Paratypes are as follows: Same 
locality data as the holotype except for 
dates, which are as follows: 15-31 May 
1965°(1'"2;"BPBM); 15 May 1966 (1° 2; 
BPBM); 15 Aug 1966 (1 2; BPBM); 30 
Jun 1967 (1 6; BPBM); 15 Aug 1967 (2 
2; BPBM, USNM); 15 Sep 1967 (1 @; 
BPBM). One <6 has been dissected (the 
structures are in an attached microvial). 

Etymology.—The species epithet, ban- 
vaneue, refers to the type locality in Laos 
and is a noun in apposition. 

Remarks.—This species is sympatric 
with C. laos but the two species are easily 
distinguished by the wing patterns. The 
wing of C. banvaneue has cell R,,, mostly 
hyaline with only the apical third dark, 
whereas in C. laos this cell is mostly dark 
with a subapical hyaline spot. A second dif- 
ference is the shape of vein R,, ;, which is 


503 


Fig. 31. 
epandrium, cerci, and surstyli, ventral view. 


Cyamops kaplanae. 20, Segments 5-7, 


straight in C. banvaneue and sinuous in C. 
laos. 


Cyamops kaplanae Baptista and Mathis, 
new species 
(Fig. 31) 


Description.—Adult ¢ length 2.1 mm; 
wing length 1.86 mm; wing width 0.6 mm. 
Adult 2 length 2.1 mm; wing length 2.2 
mm; wing width 0.8 mm. 

Head: Ocellar tubercle and vertex mostly 
polished; depressed region of frons deep 
black; antenna yellow, slightly infuscate on 
dorsal margin; face pale yellow, strongly 
microtomentose; labellum and palpus yel- 
low; face in profile angulate, sloped anter- 
oventrally from base of antenna to vibrissal 
angle, thereafter slightly receded to oral 
margin, produced portion about %4 width of 
Ist flagellomere. Chaetotaxy: Inner fronto-’ 
orbital setae slightly divergent, about half 
length of outer vertical seta; arista bearing 
10 dorsal, 3 ventral rays, 7 basalmost rays 
bifurcate; pseudovibrissae slightly divergent 
and curved dorsally; peristomal setae 7. 

Thorax: Halter mostly brown, base pale 
yellow; scutellum trapezoidal, orientation 
of scutellum moderately more elevated than 
scutum, disk flat; postpronotum and central 


504 


portion of anepimeron polished. Chaeto- 
taxy: Posterior dorsocentral seta almost 3 
times longer than anterior seta; mesonotal 
setulae moderately developed; scutellar se- 
tae 2, basal seta % length of apical seta, 
thinner. Legs mostly yellow, only hindfe- 
mur brown on apical 14; apical and sub- 
apical tarsomere of each leg brown; spine- 
like setulae 4, weakly differentiated. Wing 
mostly hyaline, with brownish area on cen- 
tral portion over cells dm, vein R,,; and 
upper portion of cell br, and apex of cell 
R,,;; cells bm and dm separate; Ist costal 
ratio 1.4; 2nd costal ratio 4.5; wing ratio 
PNG [ye 

Abdomen (Fig. 31): 6th tergite about 
same width as 7th tergite, both tergites al- 
most without setae; 6th—7th sternite with 
posterior process as in Fig. 31. Male ter- 
minalia (Fig. 31): Right surstylus large, 
subrectangular, complex in shape; left sur- 
stylus slender, length about 2 that of right 
surstylus; ejaculatory apodeme short, as 
long as 5th tergite dorsally, extended pro- 
cess thin and short, with margins subpar- 
allel. 

Female: Head: Antenna dark, median 
portion of Ist flagellomere microtomentose, 
with silver dust; produced portion of face 
larger than width of Ist flagellomere; me- 
sofacialia and gena dark brown, strongly 
microtomentose, central portion shiny; pal- 
pus brown to dark brown; femora dark ex- 
cept for yellow basal portion. 

Thorax: Wing: Hyaline 

Abdomen (abdomen not dissected): 7th 
tergite and sternite separate. 

Type material—The holotype ¢ is la- 
beled “THAILAND: S. KhaoSokNatPar. 
Rt. 401, 22.X.1993 [22 Oct 1993], EF KAP- 
LAN & A. FREIDBERG/Dissected by A. 
Baptista97 [type written/HOLOTYPE ¢ 
Cyampos kaplanae Baptista & W.N. Mathis 
USNM [red; species name, gender symbol, 
and “Baptista &’’ handwritten].’’ The ho- 
lotype is double mounted (glued to a paper 
triangle), is in excellent condition (abdo- 
men removed and dissected, structure in an 
attached microvial), and is deposited in the 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


USNM. The paratype 2? bears the same lo- 
cality data as the holotype (1 2; USNM). 

Etymology.—The species epithet, ka- 
planae, is a genitive patronym to recognize 
the collecting efforts of Ms. Fini Kaplan, 
who collected the type series of this species 
and many other specimens of interesting 
acalyptrate Diptera. 

Remarks.—The 7th tergite and sternite of 
the female abdomen are separate. Having a 
suture between the 7th tergite and the 6th— 
8th sternites is apparently a plesiomorphic 
feature. Female specimens from the Austra- 
lasian Region also have the tergite and ster- 
nite of the 7th segment separate. 


PALEARCTIC REGION 


No species of Cyamops has been de- 
scribed from this zoogeographic region. 


Cyamops sp. 4 


One specimen from Japan (Kyushu Bep- 
pa, 21 Jun 1952, P. W. Oman; USNM) has 
the following combination of characters: 
Face constricted medially; lacking mesofa- 
cial setae; pseudovibrissae aligned with per- 
istomal setae; dorsocentral setae 2. 

The only available specimen is in poor 
condition, and we defer describing this spe- 
cies until better preserved specimens be- 
come available. 


PHYLOGENETIC CONSIDERATIONS 


At the time of our revision we followed 
D. McAlpine (1989) and classified Cy- 
amops in the family Periscelididae. Roha- 
éek (1998) has questioned the placement of 
Stenomicra and Cyamops (‘‘Stenomicri- 
dae’’) with the other Periscelididae as un- 
natural, based on the “‘different types of 
male postabdomen and hypopygium’’ for 
each group. Moreover, he tentatively in- 
cluded the anthomyzid genus Echidnoce- 
phalodes Sabrosky in Periscelididae. 

We agree with Rohacek that the support- 
ing evidence for the monophyly of Peris- 
celididae is weak and needs further inves- 
tigation. That both groups have different 
types of postabdomens, however, does not 


VOLUME 102, NUMBER 3 


505 


Figs. 32-37. Wings. 32, Cyamops funkae. 33, C. nigeriensis. 34, C. fiji. 35, C. femoratus. 36, C. laos. 37, 


C. banvaneue. 


preclude the hypothesis of monophyly. The 
“naturalness” of a group does not rely 
wholly on characters of the male terminalia 
or on any one character suite in particular. 
Our study of structures of the male termin- 
alia of many Periscelididae reveals that 
these features are often so highly modified 
and derived that they obscure hypotheses of 
primary homology. 

The caplike pedicel also occurs in the 
family Neurochaetidae, and it is possible 
that Periscelididae and Neurochaetidae are 
sister groups or one family. 


ACKNOWLEDGMENTS 


This research was partially supported by 
a PEET grant from the National Science 
Foundation (PEET 952-1773). We thank 
Amnon Freidberg, Marion Kotrba, and Ste- 
phen D. Gaimari for reviewing a draft of 
this paper. We are also grateful to the 
Smithsonian Institution’s Biodiversity of 
the Guianas Program (publication 26; Vicki 
A. Funk, Director; Carol Kelloff, Coordi- 


nator) for supporting field work in Guyana. 
To Neal L. Evenhuis (BPBM) who loaned 
specimens, we express our sincere thanks. 
George L. Venable expertly produced the 
plate of wing illustrations. We are also 
grateful to Amnon Freidberg for the gen- 
erous donation of specimens that he and 
Fini Kaplan collected on Madagascar and 
in Thailand. 


LITERATURE CITED 


Baptista, A. R. and W. N. Mathis. 1994. A revision of 
New World Cyamops Melander (Diptera, Peris- 
celididae). Smithsonian Contributions to Zoology 
563: 1-28. 

. 1996. A new species of Cyamops Melander 
(Diptera: Periscelididae) from Brazil, with distri- 
butional notes on another species. Proceedings of 
the Entomological Society of Washington 98: 
245-248. 

Dallwitz, M. J., T. A. Paine, and E. J. Zurcher. 1993. 
User’s Guide to the DELTA System: a General 
System for Processing Taxonomic Descriptions. 
4th ed., 136 pp. 

Hennig, W. 1969. Neue Gattungen und Arten der Aca- 


506 


lyptratae. The Canadian Entomologist 101(6): 
589-633. 

Khoo, K. C. 1985. The Australian species of Cyamops 
Melander (Diptera: Periscelididae). Australian 
Journal of Zoology 32(1984) (2): 527-536. 

Khoo, K. C. and C. W. Sabrosky. 1989. 75. Family 
Stenomicridae, p. 551. Jn Evenhuis, N. L., ed., 
Catalog of the Diptera of the Australasian and 
Oceanian Regions. E. J. Brill and B. P. Bishop 
Museum special publication, Honolulu, 86: 1- 
1155 pp. 

McAlpine, J. E 1981. Morphology and terminology - 
adults, pp. 9-63. In McAlpine, J. EF, et al., eds., 
Manual of Nearctic Diptera. Vol. 1. Monograph 
27, Research Branch Agriculture Canada, Ottawa, 
vi. + 674 pp. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Rohaéek, J. 1998. Taxonomic limits, phylogeny and 
higher classification of Anthomyzidae (Diptera), 
with species regard to fossil record. European 
Journal of Entomology 95: 141-177. 

Sabrosky, C. W. 1958. New species and notes on North 
American acalyptrate Diptera. Entomological 
News 69: 169-176. 

. 1965. Family Anthomyzidae, pp. 819-820. In 

Stone, A., et al., eds., A Catalog of the Diptera of 

America North of Mexico. United States. Depart- 

ment of Agriculture, Agriculture Handbook No. 

276, iv + 1696 pp. 

. 1980. 66. Family Aulacigastridae, pp. 648— 

649. In Crosskey, R. W., ed., Catalogue of the 

Diptera of the Afrotropical Region. British Mu- 

seum (Natural History), London, 1437 pp. 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 507-518 


THE BIOLOGY OF DOA AMPLA (GROTE) (LEPIDOPTERA: DOIDAE) ON 
ITS HOST PLANT STILLINGIA TEXANA (EUPHORBIACEAE) 


W. EVAN BRASWELL AND JAMES R. OTT 


Department of Biology, Southwest Texas State University, San Marcos, TX 78666, 
U.S.A. (e-mail: JOO5 @academia.swt.edu): (WEB) present address: Department of Biol- 
ogy, New Mexico State University, Las Cruces, NM 88001, U.S.A.; (JRO) author for 


reprint requests 


Abstract.—We confirm Stillingia texana L. (Euphorbiaceae) as a primary host plant of 
Doa ampla Grote (Lepidoptera: Doidae), document aspects of the biology, ecology, and 
behavior of larval and adult D. ampla on S. texana, and provide initial geographic dis- 


tribution data for D. ampla in central Texas. 


Key Words: 


Doa ampla (Grote) is one of five species 
in the genus Doa (Neumoegen and Dyar 
1894). The familial placement of this group 
has a checkered history (Donahue and 
Brown 1987). The genus Doa has at times 
been placed in the Lymantriidae (Dyar 
1903, Holland 1903, Barnes and Mc- 
Dunnough 1917, Bryk 1934), Hypsidae 
(Walton 1912), Pericopidae (Schaus 1927, 
McDunnough 1938, Peterson 1948), Diop- 
tidae (Brues and Melander 1954), and Arc- 
tiidae (Franclemont 1983). The genera Doa 
and Leuculodes (which contains two spe- 
cies) currently are placed in the family Doi- 
dae within the superfamily Noctuoidea 
(Donahue and Brown 1987). According to 
Donahue and Brown, elevation of doid 
months to the family level represents an in- 
terim solution; however, Miller (1991) 
strongly supported family status. 

All known host plants of D. ampla are in 
the family Euphorbiaceae. Doa ampla pre- 
viously has been reared on Euphorbia ro- 
busta (Engelmann) in Colorado (Cockerell 
1911), E. incisa Engelmann and E. lurida 
Engelmann in Arizona, and Stillingia tex- 


gregarious behavior, aposematic coloration, group defense 


ana L. in Texas (Donahue and Brown 
1987). Doa ampla occurs from Colorado 
east to Texas, southward to Durango and 
Nuevo Le6én, Mexico, and west to Arizona 
(Donahue and Brown 1987). Aside from 
limited information on geographic distri- 
bution and host plant range, nothing is 
known of the biology and ecology of D. 
ampla. Here we confirm S. texana as a pri- 
mary host plant of D. ampla in Texas, doc- 
ument aspects of the biology, ecology, and 
behavior of larval and adult stages, and pro- 
vide distribution data for D. ampla in cen- 
tral Texas. 


METHODS 


Here we present the findings of field and 
laboratory studies conducted during the 
summers of 1996 and 1997. To confirm S. 
texana as a host plant of D. ampla (as ev- 
idenced by acceptance of S. texana for ovi- 
position and the ability of D. ampla larvae 
to complete development successfully) we 
searched S. texana plants for ovipositing fe- 
males, egg clusters, and larvae at three field 
sites. These field sites were located at Hon- 


508 


ey Creek State Natural Area (Comal Co., 
Texas), Pollard Refuge (Southwest Texas 
State University, Hays Co., Texas), and a 
privately owned site along Devils Backbone 
(Comal Co., Texas). We censused daily egg 
clusters located in the field to estimate tim- 
ing events in the life cycle, describe larval 
behavior, and document natural enemies. In 
addition, we collected females from S. tex- 
ana patches and introduced them onto pot- 
ted S. texana plants in the laboratory to 
monitor oviposition. We subsequently mon- 
itored egg clusters daily to estimate rates of 
development for each instar at 21—23°C. 
We gathered initial distribution and abun- 
dance data for D. ampla in south-central 
Texas by surveying S. texana populations 
in seven adjacent counties located along the 
eastern edge of the Edwards Plateau. Pri- 
mary state roads were used to establish 
multiple transects across each county. All 
S. texana populations on each transect that 
were accessible through a public right-of- 
way were searched for D. ampla egg clus- 
ters and larvae. We recorded the location 
(latitude, longitude, and elevation) of sites 
containing D. ampla by use of a Magellan 
GPS unit to provide site records for this ap- 
parently uncommon species (see below). 
We recorded site characteristics of occupied 
and unoccupied plant populations to de- 
scribe the habitat affinities of this herbivore. 
Initial field observations indicated that 
groups of larvae are often capable of de- 
foliating the host plant and thus must travel 
between host plants to complete develop- 
ment. To determine at which instar larvae 
are capable of dispersing to adjacent plants 
we field-tested the dispersal capabilities of 
first- through fourth-instar larvae. Five 
field-collected larvae of each instar were 
marked with fluorescent dye powder and 
placed on the ground in the center of a 1- 
m-diameter circular area bordered by a ring 
of S. texana plants. All trials were initiated 
in full sun between 1500 and 1600 to mimic 
the conditions under which late-instar lar- 
vae had been observed to move between 
plants. We then counted the number of lar- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


vae that successfully traversed the 0.5-m 
distance to a S. texana stalk after nightfall, 
using a UV lamp, and again at 24 hours. In 
total, 20 larvae of each instar were tested. 

To describe the pattern of movement of 
feeding groups within plants and the move- 
ment of individual larvae within the natal 
plant as feeding aggregations fragmented 
we marked all second-instar larvae in feed- 
ing aggregations on selected plants at the 
Honey Creek field site using fluorescent 
dye. For the duration of the second and 
third instar we then relocated these larvae 
daily during both day and night. Using D. 
ampla clutches located on a second set of 
focal plants, we then estimated the rate of 
dispersal (%) and dispersal distance of 
third-instar or older larvae from natal 
plants. On each focal plant we counted and 
marked all larvae present with a color of 
dye unique to that plant just prior to the 
onset of dispersal (i.e., immediately after 
the molt into the third instar). We then cen- 
sused each focal plant and all other S. tex- 
ana plants within 20 m of each focal plant 
for marked larvae daily during both day and 
night until larvae molted into the fifth in- 
star. Relocated larvae were re-marked after 
each molt to maintain unique marks. We 
calculated percent dispersal as the ratio of 
the number of third-, fourth-, and fifth-in- 
star larvae relocated on adjacent plants to 
the original number of third-instar larvae on 
each focal plant. The distribution of dis- 
tances moved from natal plants was used to 
construct a dispersal profile for late-instar 
larvae. We then compared the distribution 
of larval dispersal distances to the distri- 
bution of near-neighbor interplant distances 
to gauge the ability of dispersing larvae to 
locate neighboring host plants. To deter- 
mine if dispersal from the natal host plant 
is related to resource depletion, we recorded 
the percent of leaves remaining on each fo- 
cal plant at the time that larvae began to 
disperse and then used regression tech- 
niques to test for the dependency of dis- 
persal rate on resource availability per 
plant. 


VOLUME 102, NUMBER 3 


RESULTS AND DISCUSSION 


Host plant confirmation—At our field 
sites we observed female D. ampla in res- 
idence with newly deposited egg clusters on 
S. texana, and caged females readily ovi- 
posited on the leaves of S. texana in the lab. 
Following oviposition on S. texana in the 
laboratory, we subsequently reared 48 D. 
ampla larvae to pupae and reared 12 of 
those to adults. Voucher specimens are de- 
posited in the Southwest Texas State Uni- 
versity Entomology Collection. These ob- 
servations confirm S. texana as a primary 
host of D. ampla in central Texas. 

During our field studies we observed 
third- and fourth-instar D. ampla larvae 
rarely feeding on Chamaesyce acuta En- 
gelmann (Euphorbiaceae). No egg masses 
were observed on C. acuta, however, and 
no larvae found on C. acuta were relocated 
the following day on the same plant. Thus, 
this feeding appears to occur only when 
late-instar larvae are in transit between S. 
texana plants (see below). In addition to S. 
texana, two other species of Stillingia, S. 
sylvatica 1. M. Johnst. and S. treculiana 
(Muell. Arg.) I. M. Johnst., occur in Texas. 
The distribution of S. sylvatica overlaps the 
eastern and northern edges of the geograph- 
ic range of S. texana, whereas S. treculiana 
overlaps along the southwestern edge of S. 
texana’s range. We were unable to locate 
either species within the seven counties we 
surveyed. Hence, whether these species 
serve as additional hosts for D. ampla in 
regions of overlap with S. texana, as bridg- 
es to other known euphorb host plants 
whose geographic distributions occur west 
of Texas, or as both, remains unknown. 

Distribution of D. ampla in central Tex- 
as.—We collected D. ampla larvae feeding 
on S. texana in six of seven counties sur- 
veyed. Representative GPS _latitude—lon- 
gitude—elevation coordinates of collection 
sites within each county are Bandera 
(29°43.42N, 99°7.47W, 555 m); Comal 
(29°55.46N, 98°9.10W, 326 m); Gillespie 
(30°10.42N, 98°44.65W, 639 m); Hays 


509 


(29°56.13N, 98°7.24W, 393 m); Real 
(29°42.67N, 99°43.82W, 543 m); and Uval- 
de (29°45.50N, 99°31.15W, 682 m) (Fig. 1). 
In addition, D. ampla has been collected in 
Blanco County, Texas, by R. Kendall (Pe- 
demales Falls State Park; collection date, 
1973; personal communication). 

Habitat affinities —The host plant, S. tex- 
ana, is a common though patchily distrib- 
uted perennial that ranges from Oklahoma 
to Coahuila, Mexico. In Texas, the species 
is restricted to the dry calcareous soils of 
the Edwards Plateau region of south-central 
Texas, west to Val Verde Co., and north to 
Wise and Collin counties (Fig. 1). Within 
south-central Texas, we found S. texana to 
occur in two markedly different ecological 
contexts: (1) dense contiguous patches cov- 
ering up to several hectares in heavily 
grazed lowlands and (2) sparse, patchily 
distributed small populations typically lo- 
cated in Hill Country uplands associated 
with moderate to steep slopes (20—45°). In 
upland areas, S. texana is characteristically 
smaller and has fewer and shorter flowering 
stalks per root system (mean number stalks 
per plant [+ SE] upland = 4.2 + 0.41, = 
30; lowland = 25.6 + 1.9, n = 30; t = 11.4, 
P < 0.01; mean height tallest stalk per plant 
[+ SE] upland = 30.0 cm + 1.4, n = 30; 
lowland = 37.6 + 1.9, n = 30; t = 3.2, P 
< 0.01). 

We searched extensively for D. ampla on 
S. texana in both lowland and upland set- 
tings. However, we never found D. ampla 
in lowland sites and only found it in a small 
percent of upland sites examined. More- 
over, within upland habitats where D. am- 
pla was present, we rarely encountered lar- 
vae. For example, at the Honey Creek site 
(a primary study site based on the avail- 
ability of D. ampla) we found eggs or lar- 
vae on fewer than 1% of the 1,042 plants 
censused. Nighttime black light surveys 
conduced at each of our three field sites 
known to be occupied by D. ampla failed 
to attract adults during the oviposition pe- 
riod. Thus, D. ampla is relatively rare in our 
study region, appears to be a habitat spe- 


510 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


O §0 100 MILES 
i 


lentes Ihe 
hatched) in Texas. 


cialist, and is rarely encountered even with- 
in its preferred habitat. Restriction of D. 
ampla to small, sparse upland plants is 
enigmatic in that the majority of larvae in 
all but very small groups must leave the 
natal host at some point during develop- 
ment and travel to a new host plant due to 
resource limitation (Braswell 1998). There- 
fore, the absence of D. ampla in lowland 
settings, where plants are larger (i.e., more 
resources per plant and therefore reduced 
need for between-plant travel) and more 
dense (i.e., reduced risk associated with in- 
terplant travel) is counterintuitive. 

Life cycle-——Doa ampla is bivoltine in 
central Texas, exhibiting two narrowly 
overlapping generations per year. Oviposi- 
tion occurs in early to mid-May and again 


The known geographic distribution of Stillingia texana (light stippling) and Doa ampla (cross- 


in mid-July. Eggs are laid in clusters of 7 
to 231 eggs (€ = 78.5 + 7.97; n = 44) in 
parallel rows (Fig. 2). Egg clusters are 
placed on the underside of leaves on the 
upper one-fourth to one-third of the host 
plant. Occasionally, females covered a sin- 
gle leaf with eggs and moved to a second 
leaf to complete oviposition. Although field 
observations of females were rare and only 
a single female has been observed moving 
to a second leaf, the large discrepancy be- 
tween the number of eggs on the first and 
second leaf suggests that the second group 
of eggs resulted from spillover of one 
clutch and not from a second female. 
Therefore, including spillover eggs, the 
mean number of eggs per clutch is 111.5 + 
11.51; n = 31. The larvae from different 


VOLUME 102, NUMBER 3 


egg clusters generally developed as inde- 
pendent groups. These egg cluster sizes are 
much larger than the 15 to 35 eggs per clus- 
ter reported for D. ampla on host plants in 
Arizona by Donahue and Brown (1987). 

First- and second-instar D. ampla are 
gregarious (Fig. 3). As in D. dora Neu- 
moegen and Dyar (Brown 1990) and D. 
raspa Druce (Dyar 1911), all instars of D. 
ampla are boldy colored and patterned, pre- 
sumably an aposematic display associated 
with the toxicity of the host plant (Fig. 4). 
Doa ampla larvae complete five instars be- 
fore pupation (Fig. 5). In the laboratory at 
21—23°C, mean development time from egg 
hatch to pupation on cuttings of S. texana 
was 35.8 d (n = 48) and from pupation to 
adult emergence was 16.25 d (n = 12). In 
the field, mean larval development time (for 
the first generation) from first to fifth instar 
was 32.6 days; larvae spent 9.9 d (n = 8 
groups) as first instars, 8.1 d (n = 13 
groups) as second instars, 7.0 d (n = 9 
groups) as third instars, and 7.6 d (n = 10 
groups) as fourth instars. In both the labo- 
ratory and field, fifth-instar larvae leave the 
host plant to pupate. Pupation sites in the 
field are unknown. In the laboratory D. am- 
pla fifth-instar larvae spun cocoons on the 
bottom, sides, or top of their nylon screen 
cages. In contrast, D. dora is know to pu- 
pate in the debris at the base of its host 
plant (Brown 1990). The cocoon of D. am- 
pla described by Dyar (1912) is similar to 
that produced by D. dora (Brown 1990) and 
D. raspa (Dyar 1911). Individual larval 
groups followed in the field underwent ec- 
dysis relatively synchronously (all larvae in 
each group completed ecdysis within 2 
days). In early May, fifth-instar larvae ap- 
peared on new growth of the host plant pri- 
or to appearance of eggs, indicating that 
second-generation D. ampla overwinter as 
late instar larvae, pupae, or both. 

Larval feeding behavior.—First- and sec- 
ond-instar larvae feed inside a communal 
nest loosely constructed of sparse silk 
threads (Fig. 3) and feed preferentially in 
the upper one-third of the host plant. These 


511 


early-instar larvae feed by scraping the up- 
per and lower surfaces of leaves leaving be- 
hind skeletonized leaf material. The leaves 
of S. texana are defended by a well-devel- 
oped laticifer system that exudes latex 
when cut. Dussourd and Eisner (1987) 
demonstrated the defensive properties of la- 
tificer systems against herbivorous insects 
by showing that when drained of latex, for- 
merly unpalatable leaves become palatable. 
Numerous insect feeding behaviors have 
been shown to represent adaptations de- 
signed to deactivate latificer-based host 
plant defenses (Dussourd and Denno 1991). 
Latex exudate is not visible when first- and 
second-instar D. ampla larvae feed on S. 
texana. Thus the mode of feeding of early- 
instar larvae may either allow them to avoid 
the laticifer system completely or scraping 
may result in small nicks that depressurize 
the laticifer system and diminish latex flow. 
If scraping depressurizes the system, indi- 
vidual larvae in groups may avoid ingesting 
large amounts of latex via the cumulative 
effect of the group on latex flow. We hy- 
pothesize that the behaviors of leaf scraping 
and gregarious feeding of early instars 
function as adaptations to diminish the 
functionality of or circumvent the host 
plant’s laticifer system (Braswell 1998). 

Beginning with the third instar, feeding 
aggregations dissolve and larval feeding be- 
havior changes. Larvae continue to feed 
preferentially on the uppermost leaves of 
the host plant; however, third- through fifth- 
instar larvae bite through the entire leaf 
blade, often leaving behind only the main 
rib of the leaf. At this stage larvae are ex- 
posed directly to the latex exudate produced 
by the plant. On host plants with multiple 
stalks, larvae move to new stalks before 
feeding on the lower leaves of any stalk. 
Paradoxically, latex flow from injured 
leaves is highest in the uppermost portions 
of the host plant (Braswell 1998). 

Larval dispersal capabilities —When lar- 
val density is high, early-instar larvae 
quickly consume the preferred feeding area 
and, on small plants, begin to defoliate the 


512 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2-3. 2, Doa ampla egg clusters on the underside of Stillingia texana leaves. 3, Second-instar D. ampla 
larvae exhibiting gregarious feeding behavior within a loose communal nest. 


VOLUME 102, NUMBER 3 


Nn 
—_ 
W 


Fig. 4. Fifth-instar Doa ampla larvae exhibiting the aposematic color pattern typical of all instars (bright 


yellow background color with longitudinal black bands). 


entire plant. Because larvae typically avoid 
lower leaves of the plant, they must travel 
from the natal plant to another plant to con- 
tinue feeding. We observed first- and sec- 
ond-instar larvae only on host plants with 
evidence of prior gregarious feeding (i.e., 
skeletonized leaves), suggesting that early 
instars do not disperse from the natal plant. 
Third- through fifth-instar larvae, however, 
were found on host plants lacking evidence 
of prior gregarious feeding, suggesting that 
these instars do disperse. 

The results of our dispersal capability ex- 
periment confirmed these field observa- 
tions. First- and second-instar larvae were 
incapable of traveling between host plants: 
none of 40 first- and second-instar larvae 
managed to move the 0.5-m linear distance 
required to reach a host plant, and all died 
within 24 h without traveling. However, 
100% (40/40) of the third- and fourth-instar 
larvae tested traveled to a new host plant 
and initiated feeding within 8 h. Marking 


and monitoring second-instar larvae in 
feeding aggregations showed that beginning 
with the third instar, concomitant with the 
switch in feeding mode, aggregations dis- 
integrate and larvae independently move to 
new feeding positions on a plant, migrate 
to new host plants, or both. 

An average of 82% of dye-marked third- 
instar larvae left their natal plants prior to 
completing development. Most larvae set- 
tled on near-neighbor plants, and no larvae 
traveled more than 5 m from the natal plant 
(Fig. 6). Mean dispersal distance of third- 
instar or older larvae (3.6 m + 1.9, n = 
126) did not differ from the mean distance 
between nearest-neighbor plants (1.6 m + 
7.6, n = 73) in the field population of S. 
texana we examined (t-test of means with 
unequal sample sizes and unequal varianc- 
es, P > 0.05). These data suggest that 
whereas dispersing larvae may not always 
find the nearest-neighbor plant in relation to 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


20 


15 


10 


Larval Body Length (mm) 


514 
2 
a 
E 
= 
iS. 
= 
& 
= 
n 
ok 
fa) 
O 
12) 
4) 
o 
ae 
0 5 
sis, Be 


The relationship between head capsule width and body length for Doa ampla larvae, illustrating the 


five instars. Instars are represented by increasing-sized symbols. Instars exhibiting gregarious behavior are de- 


picted with shaded symbols. 


their natal plant, they are relatively adept at 
locating near-neighbor host plants. 

Third-instar larvae can markedly reduce 
the number of leaves per plant prior to the 
onset of dispersal. The percent of leaves re- 
maining per plant was inversely and non- 
linearly related to the number of third-instar 
larvae per group (R? = 0.41, P < 0.05; Fig. 
7). Moreover, the percent of larvae that dis- 
persed was negatively related to the percent 
of leaves remaining on the host plant (R? = 
0.38, P < 0.05; Fig. 8). Therefore, dispersal 
rate appears to be influenced by resource 
availability as mediated by the interaction 
of larval group size and plant size. 


Larval defensive behavior.—First and 
second instars when disturbed exhibit a 
head-flicking display. In this display, larvae 
rear up on the abdominal prolegs and thrash 
the head from side to side. The display oc- 
curs synchronously within the aggregation, 
and communication between larvae appears 
to be facilitated by the silk of the nest. Fol- 
lowing continued disturbance, larvae drop 
from the host on silken threads. In later in- 
stars the head-flicking behavior decreases 
and larvae are more likely to drop to the 
ground when disturbed. 

Predators and parasitoids.—Through two 
field seasons we observed relatively few 


VOLUME 102, NUMBER 3 


10 


Frequency 


0 100 


200 


ails 


400 500 


300 
Mean Dispersal Distance/aggregation (cm) 


Fig. 6. Mean dispersal distances/group for each of 25 groups of later-instar D. ampla larvae from their natal 


host plants under field conditions. 


acts of larval predation and no acts of larval 
parasitism. No predation or parasitism of 
eggs was observed. The only successful 
acts of predation occurred on early-instar 
larvae by ants, which collectively attacked 
and removed single larvae. The only other 
act of predation we observed was an attack 
by a lynx spider (Peucetia viricans Hentz) 
on a third-instar larva. The spider bit and 
killed the larva but never returned to con- 
sume it. We collected later-instar larvae 
from both generations that appeared to have 
been parasitized as evidenced by their slug- 
gish, unresponsive behavior. These animals 
did not feed and remained alive for months 
before dehydrating. No parasitoids emerged 
from these larvae. 


SUMMARY 


Stillingia texana is a host plant for Doa 
ampla in central Texas. Although this host 
plant is common throughout the eastern 
edge of the Edwards Plateau, often occur- 
ring in large, contiguously distributed pop- 
ulations, D. ampla is rarely encountered 
and is restricted to small, low-density 
patches of host plant occupying upland re- 
gions of moderate to steep slope. Doa am- 
pla is bivoltine in central Texas and over- 
winters as late-instar larvae, pupae, or both. 
First- and second-instar larvae feed gregar- 
iously on leaf tissue by scraping the epi- 
dermal layer. This mode of feeding circum- 
vents or reduces contact with the plant’s la- 


516 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


LOO 


75 


Percent of Leaves Remaining 
N Nn 
Nn =) 


0) 10 20 30 40 50 
Number of Third Instar Larvae per Group 


30 
2215, 
20 
1s 


10 


Percent of Larvae Which Dispersed 


0 25 50 a5 100 
Percent of Leaves Remaining 


Fig. 7-8. 7, The relationship between the number of third-instar larvae per group at the onset of dispersal 
and the percent of leaves remaining per plant (y = 90.1 — 47.6(x); R? = 0.41, P < 0.05). 8, The relationship 
between resource availability (percent of leaves remaining at the onset of dispersal) and the percent of larvae 
that disperse from their natal plant (y = 17.9 — 0.15(x); R? = 0.38, P < 0.05). 


VOLUME 102, NUMBER 3 


ticifer system. Beginning at the third instar, 
the mode of feeding switches to leaf chew- 
ing, and feeding aggregations dissolve. Lat- 
er-instar larvae continue to feed on the natal 
plant or disperse to adjacent plants. Only 
third-instar or later larvae are capable of 
dispersal, and dispersing larvae are adept at 
locating near-neighbor host plants. Dispers- 
al rate from natal plants may be governed 
by resource availability during the third lar- 
val instar, which may be mediated by feed- 
ing group size in relation to plant size. The 
ecological factors underlying dispersal from 
natal plants deserve further attention as 
there seems to be a fundamental conflict be- 
tween the clutch size deposited by females 
on plants and the amount of resources re- 
quired to support development of the 
clutch. First- and second-instar larvae ex- 
hibit group defense, and all instars are 
brightly patterned with contrasting colors. 
The apparent lack of natural enemies, along 
with the gregarious nature, defensive be- 
haviors, apparent aposematic coloration, 
and the chemical composition of the host 
plant suggest that D. ampla larvae are nox- 
ious to natural enemies and that the larval 
color pattern may function as warning col- 
oration used to deter predation. Based on 
these initial observations D. ampla popu- 
lations may be controlled by factors other 
than natural enemies. 


ACKNOWLEDGMENTS 


This research was supported by a Faculty 
Research Enhancement Grant from South- 
west Texas State University to JRO and by 
both a Ruth & Russell Strandtman Field 
Research Award and Sigma Xi Grant-In- 
Aid of Research Award to WEB. We thank 
D. Ferguson of the USDA Systematic En- 
tomology Laboratory for the identification 
of Doa ampla and J. Donahue for assistance 
with literature and classification history. 
The Southwest Texas State University Free- 
man Ranch and the Texas Parks and Wild- 
life Department provided access to field 
sites for this research. We thank J. Adams, 
J. Donahue, and an anonymous reviewer for 


517 


providing helpful editorial comments on 
this manuscript and E. Silverfine for her 
technical editing. 


LITERATURE CITED 


Barnes, W. and J. McDunnough. 1917. Check List of 
the Lepidoptera of Boreal America. Herald Press, 
Decatur, Illinois. 392 pp. 

Braswell, W. E. 1998. The Biology and Ecology of 
Gregarious Larval Behavior in Doa ampla (Lep- 
idoptera: Doidae). MS Thesis. Southwest Texas 
State University, San Marcos, Texas, USA. 

Brown, J. W. 1990. The early stages of Doa dora Neu- 
moegen and Dyar (Lepidoptera: Noctuoidea: Doi- 
dae) in Baja California, Mexico. Journal of Re- 
search on the Lepidoptera 28: 26—36. 

Brues, C. T. and A. L. Melander. 1954. Classification 
of insects. Bulletin of the Museum of Compara- 
tive Zoology 108: 238. 

Bryk, FE 1934. Lymantriidae. Jn Strand, E., ed., 
Lepidopterorum Catalogus (62). W. Junk, Berlin. 
441 pp. 

Cockerell, T. D. A. 1911. An Aleyrodes on Euphorbia, 
and its parasite (Rhynch., Hym.). Entomological 
News 22: 462—464. 

Donahue, J. P. and J. W. Brown. 1987. The family 
Doidae, pp. 534—536. Jn Stehr, E, ed., Immature 
Insects, Vol. 1. Kendall/Hunt Publishing Compa- 
ny, Dubuque, Iowa. 

Dussourd, D. E. and T. Eisner. 1987. Vein-cutting be- 
havior: Insect counterploy to the latex defense of 
plants. Science 237: 898-901. 

Dussourd, D. E. and R. EK Denno. 1991. Deactivation 
of plant defense: Correspondence between insect 
behavior and secretory canal architecture. Ecology 
72: 1383-1396. 

Dyar, H. G. 1903. A list of North American Lepidop- 
tera. Bulletin of the United States National Mu- 
seum 52: 261. 

1911. Descriptions of the larvae of some 

Mexican Lepidoptera. Proceedings of the Ento- 

mological Society of Washington 13: 227-232. 

. 1912. [untitled note describing larvae, pupae, 
and cocoon of Doa ampla]. Proceedings of the 
Entomological Society of Washington 14: 14—15. 

Franclemont, J. G. 1983. The family Arctiidae, pp. 
114-119. In Hodges, R., ed., Check List of the 
Lepidoptera of America North of Mexico. E. W. 
Classey Ltd., Wedge Entomological Research 
Foundation, London. 

Holland, W. J. 1903. The Moth Book. Doubleday, 
Page, and Company, New York. 479 pp. 

McDunnough, J. 1938. Checklist of the Lepidoptera of 
Canada and the United States of America. Part |. 
Macrolepidoptera. Memoirs of the Southern Cal- 
ifornia Academy of Science 1: 132. 

Miller, J. S. 1991. Cladistics and classification of the 


518 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Notodontidae (Lepidoptera: Noctuoidea) based on 
larval and adult morphology. Bulletin of the Amer- 
ican Museum of Natural History 204: 170-174. 

Neumoegen, B. and H. Dyar. 1894. A preliminary re- 
vision of the Bombyces of America north of Mex- 
ico. Journal of the New York Entomological So- 
ciety 2: 147-174. 


Peterson, A. 1948. Larvae of Insects, part 1. Edwards 
Brothers, Inc., Ann Arbor, Michigan. 

Schaus, W. 1927. Pericopidae, p. 563. Jn Seitz, Gross- 
Schmetterlinge der Erde Volume 6. 

Walton, W. R. 1912. Notes on certain species of flies. 
Proceedings of the Entomological Society of 
Washington 14:13—19. 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 519-532 


LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF 
NEASPILOTA APPENDICULATA FREIDBERG AND MATHIS (DIPTERA: 
TEPHRITIDAE) ON MACHAERANTHERA CANESCENS (PURSH) A. GRAY 
(ASTERACEAE) IN SOUTHERN CALIFORNIA 


RICHARD D. GOEDEN 


Department of Entomology, University of California, Riverside, CA 92521, U.S.A. 
(e-mail: rgoeden @ucrac1.ucr.edu) 


Abstract.—Neaspilota appendiculata Freidberg and Mathis is a univoltine, monopha- 
gous fruit fly (Diptera: Tephritidae) developing solely in the flower heads of Machaer- 
anthera canescens (Pursh) A. Gray (Asteraceae) belonging to the subtribe Solidagininae 
of the tribe Astereae in southern California. The egg, first-, second-, and third-instar 
larvae, and puparium are described and figured. The pedicel of the egg is completely 
circumscribed apically by different sized, semicircular to elliptical aeropyles arranged 
singly and in rows of two to three parallel to the long axis of the egg. The dorsal sensory 
organ of the first instar is well defined, but flattened, not dome-shaped, and the single 
integumental petal and stomal sense organ are fused in this instar. The lateral stelex 
sensilla are ringed basally with three, irregular, poorly developed acanthae. The anterior 
thoracic spiracle of the second instar uniquely bears eight, cuboidal papillae, reduced to 
two or three, oblong papillae in the third instar. The ventrally-toothed oral ridges number 
seven in the third instar, which compares to seven or eight in one congener, and six in 
three other congeners examined to date. The ventrally toothed appearance and arrangement 
in a vertical series of these oral ridges appears to be a generic character. The puparium 
is reniform in shape. The larvae feed mainly on the ovules and soft achenes as first and 
second instars; however, as third instars, they usually extend their feeding into the recep- 
tacle and additionally feed on sap that collects in the shallow scars. The nonfeeding 
prepuparium overwinters in a protective cell that occupies much of the excavated flower 
head and is formed of floret, pappus, ovule and achene fragments impregnated with excess 
sap and liquid feces that harden when dry. A few prepuparia pupate and emerge from 
their cells in the late summer and fall and probably overwinter as adults, but most pu- 
pariate during the next year in late-winter to early spring, and emerge as adults that 
aggregate in summer on preblossom host plants to mate and subsequently oviposit. Pter- 
omalus sp. (Hymenoptera: Pteromalidae) was reared as a solitary, larval-pupal endopar- 
asitoid, and an unidentified Eulophidae (Hymenoptera) was reared as a gregrarious en- 
doparasitoid from puparia of N. appendiculata. 


Key Words: Insecta, Neaspilota, Machaeranthera, Asteraceae, nonfrugivorous Tephri- 
tidae, biology, taxonomy of immature stages, flower-head feeding, mo- 
nophagy, seed predation, parasitoids 


Revision of the genus Neaspilota (Dip- (1986) facilitated determination of speci- 
tera: Tephritidae) by Freidberg and Mathis mens reared from California Asteraceae 


520 


(Goeden 1989) and stimulated several life- 
history studies, including those on N. virt- 
descens Quisenberry (Goeden and Headrick 
1992), N. wilsoni Blanc and Foote (Goeden 
and Headrick 1999), N. signifera (Coquil- 
lett) (Goeden 2000a), and N. aenigma 
Freidberg and Mathis (Goeden 2000b). This 
paper describes the immature stages and life 
history of a fifth species from California, 
N. appendiculata Freidberg and Mathis. 


MATERIALS AND METHODS 


The present study was based in large part 
on dissections of flower heads of Machaer- 
anthera canescens (Pursh) A. Gray collect- 
ed discontinuously during 1988-1997 
mainly from the following three locations 
in the northern section of the San Bernar- 
dino Nat. Forest, SW San Bernardino Co.: 
Onyx Peak at 2700-m elevation, Caribou 
Creek along Van Duzen Canyon Road at 
2120 m, and N shore Big Bear Lake at 2020 
m. One-liter samples of excised, immature 
and mature flower heads containing eggs, 
larvae, and puparia were transported in 
cold-chests in an air-conditioned vehicle to 
the laboratory and stored under refrigera- 
tion for subsequent dissection, photogra- 
phy, description, and measurement. Six 
eggs, six first-, 12 second-, and 13 third- 
instar larvae, and five puparia dissected 
from flower heads were preserved in 70% 
EtOH for scanning electron microscopy 
(SEM). Additional prepuparia and 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- 
methyldisilazane (HMDS), mounted on 
stubs, sputter-coated with a gold-palladium 
alloy, and studied and photographed with a 
Philips XL-30 scanning electron micro- 
scope in the Institute of Geophysics and 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Planetary Physics, 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 studies of 
longevity and sexual maturation in the in- 
sectary of the Department of Entomology, 
University of California, Riverside, at 25 + 
1°C, and 14/10 (L/D) photoperiod. Six pairs 
of virgin males and females obtained from 
emergence cages also were held in each of 
six separate, clear-plastic, petri dishes pro- 
visioned with a flattened, water-moistened 
pad of absorbant cotton spotted with honey 
(Headrick and Goeden 1994) for observa- 
tions of their courtship and copulation be- 
havior. 

Plant names used in this paper follow 
Hickman (1993) and Bremer (1994); te- 
phritid names and adult terminology follow 
Foote et al. (1993). Terminology and tele- 
graphic format used to describe the imma- 
ture stages follow Goeden (2000a, b), Goe- 
den et al. (1998), Goeden and Headrick 
(1992, 1999), Goeden and Teerink (1997; 
1998; 1999a, b), Teerink and Goeden 
(1999), and our earlier works cited therein. 
Means + SE are used throughout this paper. 
Voucher specimens of N. appendiculata im- 
mature stages, adults, and parasitoids reside 
in my research collections. 


RESULTS AND DISCUSSION 
Taxonomy 


Adult.—Neaspilota appendiculata was 
described by Freidberg and Mathis (1986, 
p. 37-39), who pictured the unpatterned 
wing (p. 72), along with drawings (p. 38) 
of the lateral aspect of the head, male right 
foretarsus, epandrium, distiphallus, epan- 
drium and cerci, aculeus and its apex en- 
larged, and spermatheca. 

Immature stages.—The egg, first-, sec- 
ond-, and third-instar larvae, and puparium 


VOLUME 102, NUMBER 3 


AccV Spot Magn Det WD Exp 
100kV 30 126x SEY 434 


Acc V Spot Magn 
100kV 3.0 _1940x 


AccV Spot Magn 
100kV30 1940x 


Det WD Exp 
SEU Allon 


Fig. 1. Egg of Neaspilota appendiculata: (A) hab- 
itus, anterior end to left; (B) pedicel showing pattern 
of aeropyles; ®&) pedicel of a different egg with its 
aeropyles. 


heretofore have not been described or fig- 
ured. 

Egg: Thirty-three eggs dissected from 
field-collected flower heads were white, 
opaque, smooth, elongate-ellipsoidal, 0.71 
+ 0.017 (range, 0.62—0.96) mm long, 0.175 


37 


+ 0.003 (range, 0.16—0.22) mm wide, 
smoothly rounded at tapered basal end (Fig. 
1A); pedicel button-like, 0.02 mm long, 
completely circumscribed apically by dif- 
ferent-sized, semicircular to elliptical, shal- 
low aeropyles, through which the spongy 
inner layers of the chorion are readily vis- 
ible; aeropyles arranged singly and in rows 
of two to three, parallel to the long axis of 
the egg (Fig. 1B, C). 

The egg of N. appendiculata is similar in 
shape to those of N. viridescens (Goeden 
and Headrick 1992) and N. wilsoni (Goeden 
and Headrick 1999), but about 20% longer 
on average than that of N. viridescens, and 
24% shorter and 27% wider on average 
than that of N. wilsoni (Goeden and Head- 
rick 1999). Moreover, N. appendiculata has 
more aeropyles that are more regularly and 
densely spaced around the pedicel apex 
than N. viridescens (Goeden and Headrick 
1992), but fewer aeropyles than N. wilsoni, 
which essentially cover the conical pedicel 
(Goeden and Headrick 1999). The eggs of 
12 species of Trupanea from California 
mainly differ from eggs of the three Neas- 
pilota spp. studied to date by having pedi- 
cels circumscribed by only one or two rows 
of aeropyles (Goeden and Teerink 1999b 
and references therein). 

First instar: White, elongate-cylindrical, 
bluntly rounded anteriorly and posteriorly 
(Fig. 2A); body segments well defined, 
nearly free of minute acanthae; gnathoce- 
phalon smooth, lacking oral ridges, with 
pair of prominent integumental petals dor- 
sad of mouth hooks (Fig. 2B-1); dorsal sen- 
sory organ well-defined, dome-shaped pa- 
pilla (Fig. 2C-1), pit sensillum at base of 
dorsal sensory organ; anterior sensory lobe 
(Fig. 2B-2, C-2) bears terminal sensory or- 
gan (Fig. 2C-3); lateral sensory organ (Fig. 
2C-4), supralateral sensory organ (Fig. 2C- 
5), and pit sensory organ (Fig. 2C-6); sto- 
mal sense organ ventrolateral of anterior 
sensory lobe (Fig. 2B-3, C-7), integumental 
petal (Fig. 2B-1) fused laterally with stomal 
sense organ (Fig. 2B-3); mouthhook biden- 
tate (Fig. 2B-4, C-8); median oral lobe lat- 


522 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


AccV SpotMagn Det WD Exp 
10.0kV 3.0 121x SE 4117 


AccY SpotMagn D 
10.0 kV 1.0 272 


Acc'V" Spot Magn - Det WD Exp f-— >——{- 10 um 
10.0 KV 1.0-°2069x ‘SE. 433.10 “-_ : 


AccY SpotMagn Det WD Exe F———{ 204m 
100kV 3.0 1034x SE 4128 


Fig. 2. First instar of Neaspilota appendiculata: (A) habitus, anterior to left; (B) gnathocephalon, ventrolat- 
eral view, l-integumental petal, 2-anterior sensory lobe, 3-stomal sense organ, 4-mouth hook, 5-median oral 


lobe; (C) gnathocephalon, frontal view, 1-dorsal sensory organ, 2-anterior sensory lobe, 3-terminal sensory organ, 
4-lateral sensory organ, 5-supralateral sensory organ, 6-pit sensory organ, 7-stomal sense organ, 8-mouthhook; 
(D) ventrolateral stellex sensillum, 1-basal acanthae; (E) anal segment, 1-rima, 2-interspiracular process; (F) 
intermediate sensory complex, 1-stelex sensillum, 2-medusoid sensillum. 


erally flattened (Fig. 2B-5); meso- and 
metathoracic and abdominal lateral spirac- 
ular complexes not seen; caudal segment 
with two stelex sensilla, dorso- and ventro- 


lateral of posterior spiracular plate, each 
stelex sensillum (Fig. 2D) basally ringed 
with three, irregular, poorly developed 
acanthae (Fig. 2D-1); posterior spiracular 


VOLUME 102, NUMBER 3 


plate bears two ovoid rimae, ca. 0.008 mm 
in length (Fig. 2E-1), and four interspira- 
cular processes, each with two to four 
branches, longest measuring ca. 0.01 mm 
(Fig. 2E-2); intermediate sensory complex 
with one stelex sensillum (Fig. 2F-1) and 
one medusoid sensillum (Fig. 2F-2). 

The first instar is similar in general hab- 
itus to that of N. viridescens (Goeden and 
Headrick 1992), N. wilsoni (Goeden and 
Headrick 1999), N. signifera (Goeden 
2000a), and N. aenigma (Goeden 2000b). 
However, unlike N. viridescens, but like N. 
wilsoni and N. aenigma, the dorsal sensory 
organ of the first instar of N. appendiculata 
is well defined (Fig. 2C-1), as is the anterior 
sensory lobe (Fig. 2B-2, C-2) and integu- 
mental petal (Fig. 2B-1). Also, the pit sen- 
sory organ (fig. 2C-6), not seen in N. viri- 
descens (Goeden and Headrick 1992) and 
presumably hidden in specimens viewed of 
N. signifera (Goeden 2000a), is present in 
N. appendiculata, as it is in N. wilsoni 
(Goeden and Headrick 1999) and N. aenig- 
ma (Goeden 2000b). A fused integumental 
petal and stomal sense organ also was re- 
ported in first instars of N. wilsoni (Goeden 
and Headrick 1999), N. signifera (Goeden 
2000a), and N. aenigma (Goeden 2000b) as 
well as Trupanea vicina (Wulp) (Goeden 
and Teerink 1999b), but these structures 
were separated in N. viridescens (Goeden 
and Headrick 1992) 

Having two stelex sensilla dorso- and 
ventrolaterad of each posterior spiracular 
plate in the first instar of N. appendiculata 
(Fig. 2D) and N. aenigma (Goeden 2000b) 
agreed with the four such sensilla reported 
to ring the caudal segment of N. wilsoni 
(Goeden and Headrick 1999), but not the 
10 sensilla reported to ring the caudal seg- 
ment of N. viridescens (Goeden and Head- 
rick 1992). The last number is probably er- 
roneous, as discussed by Goeden (2000b). 
Lateral stelex sensilla on the caudal seg- 
ment that are basally ringed with acanthae 
(Fig. 2D-1) first were reported in N. wilsoni 
(Goeden and Headrick 1999), where the up- 
right acanthae among them number one to 


523 


three and are pointed, and in N. aenigma 
(Goeden 2000b), where the upright acan- 
thus is solitary and rounded apically. 

Second instar: White, elongate-cylindri- 
cal, rounded anteriorly, truncated dorsopos- 
teriorly (Fig 3A), body segments well de- 
fined, circumscribed by only a few minute 
acanthae; dorsal sensory organ well-defined 
(Fig. 3B-1, C-1), with basally associated 
pore sensilla (Fig. 3C-2); anterior sensory 
lobe (Fig. 3B-2, C-3) with terminal sensory 
organ (Fig. 3B-3, C-4), lateral sensory or- 
gan (Fig. 3C-5), supralateral sensory organ 
(Fig. 3C-6), and pit sensory organ (Fig. 3C- 
7); stomal sense organ ventrolaterad of an- 
terior sensory lobe (Fig. 3B-4, C-8); mouth- 
hook bidentate (Fig. 3B-5); median oral 
lobe laterally compressed (Fig. 3B-6), ven- 
trally transversely divided (not shown); sev- 
en papilliform, integumental petals dorsal 
to each mouthhook (Fig. 3B-7, C-9); six 
oral ridges toothed ventrally, lateral to oral 
cavity (Fig. 3B-8); pore sensilla circum- 
scribe gnathocephalon posterior to oral 
ridges (Fig. 3B-9); prothorax circumscribed 
anteriorly by minute acanthae (Fig. 3B-10); 
anterior thoracic spiracle with eight cuboi- 
dal papillae (Fig. 3D); lateral spiracular 
complexes not seen; caudal segment with 
two stelex sensilla dorsolaterad and ventro- 
laterad of posterior spiracular plate (not 
shown); posterior spiracular plate bears 
three ovoid rimae, ca. 0.02 mm long, and 
four interspiracular processes, each with 
four, simple or forked branches, longest 
measuring 0.012 mm; intermediate sensory 
complex with a stelex sensillum and a me- 
dusoid sensillum. 

The habitus of the second instar of N. 
appendiculata (Fig. 3A) is more like N. wil- 
soni (Goeden and Headrick 1999), N. sig- 
nifera (Goeden 1999a), and N. aenigma 
(Goeden 2000b) than the barrel-shaped sec- 
ond instar of N. viridescens (Goeden and 
Headrick 1992). The dorsal sensory organ 
of N. appendiculata is well defined in the 
second instar (Fig. 3C-1), as with N. sig- 
nifera (Goeden 1999a), but is not well de- 
fined in N. viridescens (Goeden and Head- 


524 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


AccV SpotMagn Det WD Exp H——{ 200um 


AccV Spot Magn 
10.0 kV 10 79x 6 Ds 280x 


10.0 kV 1.0 


Fig: 3: 


Spot Magn Det 
STOOKV'3.0 321 7x SE  41.3/6 


WD Exp 


Second instar of Neaspilota appendiculata: (A) habitus, anterior to left; (B) gnathocephalon, ventro- 


lateral view, 1-dorsal sensory organ, 2-anterior sensory lobe, 3-terminal sensory organ, 4-stomal sense organ, 
5-mouthhook, 6-median oral lobe, 7-integumental petal, 8-oral ridge, 9-pore sensillum, 10-minute acanthae; (C) 
anterior sensory lobe, 1-dorsal sensory organ, 2-basilateral, pore sensillum, 3-anterior sensory lobe, 4-terminal 
sensory organ, 5-lateral sensory organ, 6-supralateral sensory organ, 7-pit sensory organ, 8-stomal sense organ, 


9-integumental petal; (D) anterior spiracle. 


rick 1992), N. wilsoni (Goeden and Head- 
rick 1999), or N. aenigma (Goeden 2000b). 
The integumental petals of the second in- 
stars of all five species are papilliform and 
seven in number above each mouthhook in 
N. appendiculata (Fig. 3B-7), but four oc- 
cur in N. signifera (Goeden 2000a), six in 
N. viridescens (Goeden and Headrick 
1992), seven in N. wilsoni (Goeden and 
Headrick 1999), and eight occur in N. 
aenigma (Goeden 2000b); whereas, in the 
first instars of all five congeners examined 
these structures are broad, flattened, and 
paired (Fig. 2B-1; Goeden and Headrick 
1992, 1999; Goeden 2000a, b). A clear dif- 
ference in N. appendiculata is the eight pa- 
pillae on the anterior spiracle of the second 


instar (Fig. 3D), compared to three to four 
papillae in second instars of N. viridescens 
(Goeden and Headrick 1992), N. wilsoni 
(Goeden and Headrick 1999), N. signifera 
(Goeden 2000a), and N. aenigma (Goeden 
2000b). Only the third-instar of Paracantha 
gentilis Hering with seven to eight papillae 
on the anterior spiracle (Headrick and Goe- 
den 1990) approached this number among 
the 34 species of tephritid larvae that we 
have described in similar detail to date. Fi- 
nally, the interspiracular processes each 
bear four branches, not one to four branches 
like N. aenigma (Goeden 2000b), nor two 
to four branches like N. signifera (Goeden 
1999a), nor five to nine branches like those 
of N. viridescens (Goeden and Headrick 


VOLUME 102, NUMBER 3 


1992), nor two to six branches like those of 
N. wilsoni (Goeden and Headrick 1999). 
Third instar: Pale yellow, with posterior 
spiracular plate dark brown to black, elon- 
gate-cylindrical, tapering anteriorly; poste- 
rior spiracular plate on caudal segment flat- 
tened and upturned dorsally ca. 60° (Fig. 
4A), minute acanthae circumscribe anterior 
thirds of thoracic and abdominal segments 
(Fig. 4B-1, C-1, D-1); gnathocephalon con- 
ical (Fig. 4B); dorsal sensory organ a well- 
defined, circular, flattened pad (Fig. 4B-2); 
anterior sensory lobe (Fig. 4B-3) bears ter- 
minal sensory organ (Fig. 4B-4), lateral 
sensory organ (Fig. 4B-5), supralateral sen- 
sory organ (Fig. 4B-6), and pit sensory or- 
gan (not shown); six, papilliform, integu- 
mental petals above each mouthhook (Fig. 
4B-7); seven oral ridges toothed ventrally 
lateral to oral cavity (Fig. 4B-8); stomal 
sense organ ventrolateral of anterior sen- 
sory lobe (Fig. 4B-9); mouthhook tridentate 
(Fig. 4B-10); median oral lobe laterally flat- 
tened (Fig. 4B-11); prothorax circum- 
scribed by minute acanthae (Fig. 4B-1); 
verruciform sensilla circumscribe prothorax 
posteriorad of minute acanthae (Fig. 4B- 
12); anterior thoracic spiracle on posterior 
margin of prothorax bears two or three ob- 
long papillae; mesothoracic lateral spiracu- 
lar complex consisting of spiracle (Fig. 4C- 
2) and five verruciform sensilla (Fig. 4C-3), 
two above and three below the spiracle; 
metathoracic lateral spiracular complex 
consisting of five verruciform sensilla (Fig. 
4C-4), one above and four below the spi- 
racle (Fig. 4C-5); abdominal lateral spirac- 
ular complex consists of a spiracle (Fig. 
4D-1) anterior to a verruciform sensillum 
(Fig. 4D-2), and two or three other verru- 
ciform sensilla, one above (not shown in 
Fig. 4D) and one (Fig. 4D-2) or two (un- 
published data) below the spiracle; caudal 
segment circumscribed by minute acanthae 
(Fig. 4E-1); each posterior spiracular plate 
bears three ovoid rimae, ca. 0.03 mm in 
length (Fig. 4E-2), and three-four interspi- 
racular processes (Fig. 4E-3), each with 
four, simple, pointed or forked branches, 


525 


longest branch measuring 0.015 mm; stelex 
sensilla (Fig. 4F) dorsolaterad, laterad, and 
ventrolaterad of posterior spiracular plate 
(Fig. 4E); each of the eight stelex sensilla 
surrounding the posterior spiracular plate in 
turn ringed by four to six, conical minute 
acanthae (4F-1); intermediate sensory com- 
plex with a medusoid sensillum and a stelex 
sensillum. 

The habitus of the third instar of N. ap- 
pendiculata generally is like that reported 
for N. viridescens (Goeden and Headrick 
1992), N. wilsoni (Goeden and Headrick 
1999), N. signifera (Goeden 2000a), and N. 
aenigma (Goeden 2000b). Like N. signifera 
(Goeden 2000a), the anterior part of each 
body segment of N. appendiculata is cir- 
cumscribed by minute acanthae. Whereas, 
in N. aenigma the anteriors, pleura, and 
posteriors of each segment are thus circum- 
scribed (Goeden 2000b); in N. wilsoni, all 
intersegmental areas and all abdominal seg- 
ments except the pleura are so circum- 
scribed (Goeden and Headrick 1999); and 
in N. viridescens, the intersegmental areas 
are free of acanthae (Goeden and Headrick 
1992). Unlike N. viridescens (Goeden and 
Headrick 1992) and N. wilsoni (Goeden and 
Headrick 1999), but like N. signifera (Goe- 
den 2000a) and N. aenigma (Goeden 
2000b), the dorsal sensory organ is well de- 
fined, but flattened and not dome-shaped in 
the third instar of N. appendiculata (Fig. 
4B-2). In the second instar of N. wilsoni 
(Goeden and Headrick 1999), N. signifera 
(Goeden 2000a), and N. appendiculata 
(Fig. 3B-2), the dorsal sensory organ is 
both prominent and dome-shaped, as it is in 
the first instar of all congeners except N. 
signifera, where it was hidden in my spec- 
imens and could not be examined for com- 
parison (Goeden 2000a). 

Additional similarities involved the inte- 
gumental petals in the third instars of all 
five congeners examined to date, all of 
which are papilliform and arranged in a 
double row above each mouth hook (Goe- 
den and Headrick 1992, 1999; Goeden 
2000a, b; Fig. 4B-7). The stomal sense or- 


526 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


AccV. Spot Magn Det Acc.V SpotMagn Det WD Exp 4H 


10.0kV10 17x SE ; T0.0kV 1.0 569x SE 403 4 


. . K A bt 
Acc.V SpotMagn Det WD EX = - A A Acc.V Spot Magh Det 
10.0KV1.0 17x SE 40.1 41 a -: Sees 


Sa 
BS Exp 
as 

Fig. 4. Third instar of Neaspilota appendiculata: (A) habitus, anterior to left; (B) gnathocephalon, lateral 
view, l-minute acanthae, 2-dorsal sensory organ, 3-anterior sensory lobe, 4-terminal sensory organ, 5-lateral 
sensory organ, 6-supralateral sensory organ, 7-integumental petal, 8-oral ridge, 9-stomal sense organ, 10-mouth- 
hook, 11-median oral lobe, 12-verruciform sensillum; (C) meso- (left) and metathoracic (right), lateral spiracular 
complexes; |-minute acanthae, 2-spiracle on mesothorax, 3-verruciform sensillum of mesothorax, 4-verruciform 

sensillum on metathorax, 5-spiracle on metathorax; (D) abdominal lateral spiracular complex; 1-spiracle, 2- 


verruciform sensillum; (E) posterior spiracular plate; 1-minute acanthae, 2-rima, 3-interspiracular process; (F) 
stelex sensillum; |-basal, conical, minute acanthus. 


en < ete AccV. Spot Magh Det W. 
ine 8S a ai. Vote — 5 » AN 


0,.0-KV.1.0.- 2738x__SE—-A 


VOLUME 102, NUMBER 3 


gan of the third instar of N. appendiculata 
(Fig. 4B-9) was not seen clearly enough to 
allow comparison with other species; how- 
ever, the third instars of all five species of 
Neaspilota examined to date have oral ridg- 
es with dentate ventral margins character- 
istically arranged in vertical series ventro- 
laterad of the dorsal sensory organ and lat- 
erad of the oral cavity. Similar oral ridges 
also were described in the second instars of 
N. viridescens (Goeden and Headrick 
1992), N. wilsoni (Goeden and Headrick 
1999), and N. signifera (Goeden 2000a). 
The oral ridges number seven or eight in 
the third instar of N. aenigma (Goeden 
2000b), seven in N. appendiculata (Fig. 
4B-8), but six in the second and third in- 
stars of the other three congeners examined 
to date. The appearance and arrangement of 
these oral ridge appears to be a generic 
character; however, the present study con- 
firms that the oral ridges vary in number 
among Neaspilota species. The third instars 
of Trupanea imperfecta (Coquillett), 7. jo- 
nesi Curran, T. nigricornis (Coquillett), T. 
pseudovicina (Hering), T. signata Foote, 
and 7. wheeleri Curran also bear serrated 
oral ridges (Goeden and Teerink 1997, 
1998, 1999a; Goeden et al. 1998; Knio et 
al. 1996; Teerink and Goeden 1999), but 
these oral ridges appear to be fewer in num- 
ber, and are not arranged in a more or less 
regular, vertical row laterad to the oral cav- 
ity, as in Neaspilota. 

The mouth hooks of the third instars of 
N. appendiculata, N. aenigma, N. signifera, 
and N. viridescens are tridentate (Goeden 
and Headrick 1992; Goeden 2000a, b); 
whereas, those of the third instar of N. wil- 
soni are bidentate (Goeden and Headrick 
1999). Such interspecific differences in 
dentation are supported by our findings that 
the mouth hooks of third-instar Trupanea 
vicina are bidentate; whereas, those of 12 
other congeners examined from California 
are tridentate (Goeden and Teerink 2000b 
and citations therein). 

The number and appearance of the stelex 
sensilla surrounding the posterior spiracular 


527 


plate differ among the Neaspilota species 
examined to date. These number only four 
in the first instars of N. wilsoni (Goeden and 
Headrick 1999). N. aenigma (Goeden 
2000b), and N. appendiculata, but unfor- 
tunately were not observed with N. signi- 
fera (Goeden 2000a). This count of stelex 
sensilla remains at four in the second instars 
of N. aenigma (Goeden 2000b) and N. ap- 
pendiculata, but increases to six in third in- 
stars of N. wilsoni (Goeden and Headrick 
1999), N. aenigma (Goeden 2000b), and N. 
appendiculata. These stelex sensilla also 
show inter-instar (intraspecific) and inter- 
specific differences in the incidence and ap- 
pearance of the minute acanthae that ring 
them basally, but this was not recognized, 
studied or recorded by my coworkers and 
me until recently (Goeden 2000b and above 
description). 

Puparium: Mostly pale yellow, with pos- 
terior two-three segments grayish to black- 
ened, reniform, and smoothly rounded at 
both ends (Fig. 5A); anterior end bears the 
invagination scar (Fig. 5B-1) and anterior 
thoracic spiracles (Fig. 5B-2); caudal seg- 
ment circumscribed by minute acanthae; 
three stelex sensilla, dorsolaterad, laterad, 
and ventrolaterad of posterior spiracular 
plates; posterior spiracular plate bears three 
broadly elliptical rimae (Fig. 5C-1), and 
four interspiracular processes, each with 
three to four branches (Fig. 5C-2); inter- 
mediate sensory complex with a medusoid 
sensillum and a stelex sensillum. Eleven 
puparia averaged 3.5 + 0.08 (range, 3.13- 
3.84) mm in length; 1.51 + 0.03 (range, 
1.42—1.71) mm in width. 


DISTRIBUTION AND Hosts 


Freidberg and Mathis (1986) described 
the distribution of N. appendiculata as, 
‘““Widespread in western United States be- 
tween 32° and 46° north latitude and be- 
tween 104° and 118° west longitude.”’ Foote 
et al. (1993) mapped the distribution to in- 
clude two or more collection records each 
from Arizona, Nevada, New Mexico, Utah, 
and Wyoming, along with additional single 


528 


Acc V Spot Magn Det WD Exp -———————_+| 1mm A 


100kV 10 23x E 39:7. 27 


nies 
7% 
4 


x 


(Ae yg PY id 
Me AAA hee) 


Fig. 5. Puparium of Neaspilota appendiculata: (A) 
habitus, anterior to left; (B) anterior end, 1-invagina- 
tion scar, 2-anterior thoracic spiracle; (C) caudal seg- 
ment, l-rima, 2-interspiracular process. 


collection records from California and Ida- 
ho, and a “‘state record only”? from Colo- 
rado. Besides our three study sites, two oth- 
er collection sites in California were along 
Pine Creek Road at 1774-m elevation, Inyo 
Nat. Forest, Inyo Co., 10.x.1990, and 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Mountain Pass, S of Interstate Highway I- 
15 at 1369 m, E. San Bernardino Co., 
GsA1998; 

The only reported and confirmed host 
plant of N. appendiculata is Machaeranth- 
era canescens (Goeden 1989), belonging to 
the subtribe Solidagininae of the tribe As- 
tereae in the family Asteraceae (Bremer 
1994). This herbaceous, annual to short- 
lived perennial, host-plant species has at 
least five distinct varieties and itself is 
widely distributed in many habitats 
throughout western United States and into 
adjacent Canada and Mexico (Hickman 
1993, Shreve and Wiggens 1964). There- 
fore, N. appendiculata provides still another 
apparent example of a true monophage re- 
ported among the nonfrugivorous Tephriti- 
dae (Headrick and Goeden 1998). 


BIOLOGY 


Egg.—In 16, mostly closed, preblossom, 
immature flower heads of M. canescens, 35 
eggs were inserted pedicel-last, six (17%) 
with their long axis at a slight angle to the 
receptacle, and 29 (83%) with their long 
axis perpendicular to the receptacle. These 
16 flower heads each contained an average 
of 2.2 + 0.4 (range, 1—7) eggs. Three (9%) 
eggs were inserted between the phyllaries 
and outer ovules and 32 (91%) within sin- 
gle florets; 12 (34%) of the 35 eggs were 
inserted into corollas of peripheral florets 
and 20 (66%) into corollas of central flo- 
rets. The diameters of the receptacles of 
these flower heads containing eggs aver- 
aged 2.4 + 0.14 (range, 1.58—3.7) mm. 

Larva.—Upon eclosion, the first instars 
usually tunneled into an ovule, or into a co- 
rolla before entering the ovule to which it 
was basally attached. Single first instars 
were found feeding within each of six, 
closed, preblossom or open flower heads. 
The receptacles of these flower heads av- 
eraged 2.4 + 0.2 (range, 1.6—2.9) mm in 
diameter. An average of 3.3 + 1.0 (range, 
2-8) ovules of soft achenes was damaged 
in these six heads as the first instar tun- 
nelled into the layer of ovules or soft 


VOLUME 102, NUMBER 3 


¥ 
4 
' 

4 
iv 

' 
Ny) 
\i 
\" 


7 


Fig. 6. 


529 


Life stages of Neaspilota appendiculata in Machaeranthera canescens: (A) second instar feeding on 


soft achenes in open flower head, (B) third instar feeding on soft achenes in open flower head, (C) prepuparium 
in center of flower head, (D) puparium of nondiapausing individual formed in flower head in summer, (E) 
puparium formed in spring by overwintered prepuparium in flower head, (F) adult male. Lines = 1 mm. 


achenes and parallel to and above the re- 
ceptacle. No receptacles within these six in- 
fested flower heads were abraded or pitted 
by feeding. Based on 54 (range, 20-75) as 
the average total number of ovules and 
achenes, respectively, counted in 29 pre- 
blossom to postblossom flower heads, about 
6% (range, 4—15%) of the ovules in the six 
infested, preblossom flower heads were 
damaged by first instars. 

Second instars continued feeding on 
ovules in preblossom flower heads or in soft 


achenes in open, blossom and post blossom, 
flower heads (Fig. 6A). All fed within a se- 
ries of adjacent ovules/soft achenes with 
their bodies horizontal to and their mouth- 
parts directed towards the receptacles, but 
always well above the receptacles. Recep- 
tacles of 14 flower heads containing second 
instars were not fed upon and averaged 2.94 
+ 0.29 (range, 1.99—5.7) mm in diameter. 
These flower heads each contained an av- 
erage of 1.2 + 0.2 (range, 1-3) larvae that 
had destroyed an average of 10.3 + 2.1 


530 


(range, 2-31) ovules/soft achenes, or as cal- 
culated for the preceding instar, about 19% 
(range, 4—-57%) of the average total of 54 
ovules/soft achenes per flower head. 

Third instars initially continued to feed 
mainly on soft achenes in post blossom 
flower heads; however, prior to pupariation, 
and usually before all the achenes were 
damaged, they proceeded to tunnel into the 
center of the receptacle (Fig. 6B). Sixty-five 
flower heads that averaged 3.09 + 0.14 
(range, 1.71—7.19) mm in diameter each 
contained a single third instar, so intraspe- 
cific mortality occurs in heads containing 
more than one larva. An average of 51 + 
7.1 (range, 24-63) of the soft achenes 
therein were damaged, or about 94% 
(range, 44—100%) of the average total of 54 
ovules/soft achenes per flower head. These 
percentages of seed predation per larva per 
flower head are on the high side among flo- 
rivorous tephritids studied by us to date 
(Headrick and Goeden 1998); this seed pre- 
dation was exceeded only by gregarious flo- 
rivorous species like Trupanea conjuncta 
(Adams) (Goeden 1987) and T. pseudovi- 
cina Hering (Goeden and Teerink 1998) or 
by species with large larvae that develop in 
immature or small flower heads like Para- 
cantha cultaris (Coquillett) (Cavender and 
Goeden 1984) and Xenochaeta dichromata 
Snow (Goeden and Teerink 1997). 

Third instars in flower heads fed with 
their long axes oriented perpendicular to 
and mouthparts directed towards the recep- 
tacles (Fig. 6B). Ninety percent of the third 
instars in the 65 infested heads scored or 
pitted the receptacles and thus presumma- 
bly supplemented their diet with sap. Goe- 
den and Headrick (1992, 1999) described 
and discussed this similar type of feeding 
by N. viridescens and N. wilsoni. And, as 
also reported for both of these congeners 
(Goeden and Headrick 1992, 1999), most 
third instars became surrounded for about 
90% their lengths by cells, which occupied 
most of the interior of the flower heads and 
consisted of ovule-, achene-, chaff-, pap- 
pus-, and corolla-fragments cemented to- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


gether by liquid feces and sap that hardened 
when dry (Fig. 6C). These protective cells 
were slightly larger than the mature larva, 
externally incorporated the outer walls of 
achenes and the few uneaten achenes, and 
were blackened and smooth inside. Upon 
completing feeding and cell construction, 
the third instars oriented with their anterior 
ends towards the receptacles, retracted their 
mouthparts, and formed prepuparia (Head- 
rick and Goeden 1998). Most individuals 
overwintered in diapause as prepuparia 
(Fig. 6C) (Goeden and Headrick 1992, 
1999; Headrick and Goeden 1998), but a 
few individuals pupariated early and 
emerged in late-summer and fall (August— 
September) (Fig. 6D). Prior to pupariation 
the prepuparia reversed their orientation 
within their cells and turned 180° such that 
their heads were directed away from the re- 
ceptacles (Fig. 6D, E). 

Pupa.—The receptacles of 11, overwin- 
tered, postblossom flower heads, each also 
containing a single puparium (Fig. 6E), av- 
eraged 3.50 + 0.08 (range, 3.13—3.84) mm 
in diameter. 

Adult.—Adults emerged from overwin- 
tered, mature flower heads, and were long- 
lived under insectary conditions, as 24 un- 
mated males (Fig. 6F) averaged 64 + 9 
(range, 10-177) days, and 14 virgin fe- 
males averaged 73 + 12 (range, 10—133) 
days. Such lengthy longevities compare fa- 
vorably with average adult longevities re- 
ported for adults of N. viridescens (Goeden 
and Headrick 1992), N. wilsoni (Goeden 
and Headrick 1999), N. signifera (Goeden 
1999a), and N. aenigma (Goeden 1999b). 

The premating and mating behaviors of 
N. appendiculata were not studied in the 
field, but were observed in petri dish arenas 
found to be so useful with many other non- 
frugivorous tephritid species (Headrick and 
Goeden 1994). Premating behaviors ob- 
served with paired N. appendiculata (n) 
were brief “‘kissings’”” (n = 2), side-step- 
ping by males while tracking females (n = 
4) (Headrick and Goeden 1994), and rapid 
wing hamation, sometimes combined with 


VOLUME 102, NUMBER 3 


lofting about 20° by both sexes (n = 6) 
(Headrick and Goeden 1994). Two matings 
were observed that began during late after- 
noon in twilight involving different pairs of 
flies that lasted 225 and 345 min. These 
compared to average durations of 190 min 
reported for N. aenigma (Goeden 2000b), 
235 min. reported for N. wilsoni (Goeden 
and Headrick 1999), 238 min. reported for 
N. signifera (Goeden 2000a), and 5.3 h re- 
ported for WN. viridescens (Goeden and 
Headrick 1992). No post-copulatory behav- 
ior reminescent of the mate guarding ob- 
served with N. signifera (Goeden 2000a), 
Dioxyna sororcula (Wiedemann) (Headrick 
et al. 1996), and Euaresta stigmatica Co- 
quillett (Headrick et al. 1995) was observed 
with N. appendiculata. Postcopulatory be- 
havior by N. appendiculata mainly consist- 
ed of storing of genitalia by males and 
cleaning and grooming by both sexes 
(Headrick and Goeden 1994) 

Seasonal history.—The life cycle of N. 
appendiculata in southern California fol- 
lows an aggregative pattern (Headrick and 
Goeden 1994, 1998) in which the prepu- 
parium is the principal overwintering stage. 
Come late spring (April-May), overwin- 
tered prepuparia reverse their orientation in 
their cells in flower heads on shoots of dead 
host plants and pupariate. Adults emerge 
during May and June and aggregate on pre- 
blossom shoots of M. canescens to mate. 
Females oviposit in the small, newly- 
formed, closed, preblossom flower heads in 
June and July and larvae feed until fully 
grown, then enter diapause in the late sum- 
mer and early fall (August—October). There 
is a single generation per year on their sole 
host plant, although as mentioned above, a 
few adults emerge in August-September, 
perhaps to produce a partial generation on 
late-flowering plants, or to overwinter as 
long-lived adults. 

Natural enemies.—Two males and two 
females of Pteromalus sp. (Hymenoptera: 
Pteromalidae) were reared from separate 
puparia of N. appendiculata as solitary, lar- 
val-pupal endoparasitoids. Seven, 15, and 


531 


17 individuals of an unidentified Eulophi- 
dae (Hymenoptera) respectively were 
reared from three puparia as gregarious en- 
doparasitoids. 


ACKNOWLEDGMENTS 


I thank Andrew C. Sanders, Curator of 
the Herbarium, Department of Botany and 
Plant Sciences, University of California, 
Riverside, for identifications of plants men- 
tioned in this paper. Krassimer Bozhilov in 
the Institute of Geophysics and Planetary 
Physics, University of California, River- 
side, greatly facilitated my scanning elec- 
tron microscopy. The parasitoids were iden- 
tified by Harry E. Andersen, Huntington 
Beach, California. I also am grateful to Jeff 
Teerink and Kristine Gilbert for technical 
assistance and to Louie Blanc, Jeff Teerink, 
and David Headrick for their helpful com- 
ments on 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., 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. 

Freidberg, A and W. N. Mathis. 1986. Studies of Ter- 
elliinae (Diptera: Tephritidae): A revision of the 
genus Neaspilota Osten Sacken. Smithsonian 
Contributions to Zoology 439: 1—75. 

Goeden, R. D. 1987. Life history of Trupanea con- 
jJuncta (Adams) on Trixis californica Kellogg in 
southern California (Diptera: Tephritidae). The 
Pan-Pacific Entomologist 63: 284-291. 

. 1989. Host plants of Neaspilota in California 

(Diptera: Tephritidae). Proceedings of the Ento- 

mological Society of Washington 91: 164-168. 

. 2000a. Life history and description of im- 

mature stages of Neaspilota signifera (Coquillett) 

(Diptera: Tephritidae) on Hemizonia pungens 

(Hooker and Arnott) Torrey and A. Gray (Aster- 

aceae) in southern California. Proceedings of the 

Entomological Society of Washington 102: 69— 

81. 


. 2000b. Life history and description of im- 
mature stages of Neaspilota aenigma Freidberg 


Nn 
Ww 
i) 


and Mathis (Diptera: Tephritidae) on Erigeron 
divergens Torrey and Gray (Asteraceae) in south- 
ern California. Proceedings of the Entomological 
Society of Washington 102: 384-397. 

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. 

. 1999. Life history and description of imma- 
ture stages of Neaspilota wilsoni Blanc and Foote 
(Diptera: Tephritidae) on Hazardia squarrosa 
(Hooker and Arnott) E. Greene (Asteraceae). Pro- 
ceedings of the Entomological Society of Wash- 
ington 101: 897—909. 

Goeden, R. D. and J. A. Teerink. 1997. Life history 
and description of immature stages of Xenochaeta 
dichromata Snow (Diptera: Tephritidae) on Hier- 
acium albiflorum Hooker in central and southern 
California. Proceedings of the Entomological So- 
ciety of Washington 99: 597-607. 

. 1998. Life history and description of imma- 

ture stages of Trupanea pseudovicinia Hering 

(Diptera: Tephritidae) on Porophyllum gracile 

Bentham (Asteraceae) in southern California. Pro- 

ceedings of the Entomological Society of Wash- 

ington 100: 361—372. 

. 1999a. Life history and description of 1m- 

mature stages of Trupanea wheeleri Curran (Dip- 

tera: Tephritidae) on Asteraceae in southern Cal- 
ifornia. Proceedings of the Entomological Society 

of Washington 101: 414—427. 

. 1999b. Life history and description of im- 
mature stages of T7rupanea vicina (Wulp) (Diptera: 
Tephritidae) on wild and cultivated Asteraceae in 
southern California. Proceedings of the Entomo- 
logical Society of Washington 101: 742-755. 

Goeden, R. D., J. A. Teerink, and D. H. Headrick. 
1998. Life history and description of immature 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


stages of Trupanea jonesi Curran (Diptera: Te- 
phritidae) on native Asteraceae in southern Cali- 
fornia. Proceedings of the Entomological Society 
of Washington 100: 126-140. 

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. 

1994. Reproductive behavior of California 

fruit flies and the classification and evolution of 

Tephritidae (Diptera) mating systems. Studia Dip- 

terologica 1(2): 194-252. 

. 1998. The biology of nonfrugivous tephritid 
fruit flies. Annual Review of Entomology 43: 
217-241. 

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. 

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. 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. 

Shreve, F and I. L. Wiggens. 1964. Vegetation and 
Flora of the Sonoran Desert. Vol. 2. Stanford Uni- 
versity Press, Stanford, California. 

Teerink, J. A. and R. D. Goeden. 1999. Description of 
the immature stages of Trupanea imperfecta (Co- 
quillett). Proceedings of the Entomological Soci- 
ety of Washington 101: 225—232. 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 533-541 


MORPHOMETRIC VARIATION AMONG POPULATIONS OF 
AMBRYSUS MORMON MONTANDON (HETEROPTERA: NAUCORIDAE) 


ROBERT W. SITES AND MICHAEL R. WILLIG 


(RWS) Enns Entomology Museum, Department of Entomology, University of Missouri, 
Columbia, Missouri 65211 (e-mail: bugs@showme.missouri.edu); (MRW) Program in 
Ecology and Conservation Biology, Department of Biological Sciences and The Museum, 
Texas Tech University, Lubbock, Texas 79409-3131 and National Center for Ecological 
Analysis and Synthesis, University of California, 735 State Street, Suite 300, Santa Bar- 
bara, California 93101-5504 (e-mail: cmmrw @ttacs.ttu.edu) 


Abstract.—Morphometric variation with respect to 15 mensural characters was assessed 
for adult specimens from 13 populations of Ambrysus mormon Montandon in the United 
States. This provides a context from which to assess the degree of divergence of an 
isolated population (Ash Warm Springs, Nevada), which possesses discrete-state charac- 
teristics that differ markedly from those of other populations of the species. A multivariate 
analysis of variance revealed that interpopulational differences were dependent on sex, a 
finding corroborated by the univariate perspective provided by Bonferroni’s sequential 
adjustment that additionally identified lengths of body, protibia, and protarsus as contrib- 
uting to the interaction between population and sex. Moreover, all pairwise comparisons 
of the 13 populations were significant for males as well as for females (F-tests from 
discriminant function analysis). Indeed, the pattern of morphometric variation among pop- 
ulations was similar in males and females but was not a consequence of differentiation 
by geographic distance (Mantel analyses). Interindividual variation primarily was due to 
differences in size, as the first axis from principal components analysis accounted for 
92.2% of the total variation. Three additional axes represented shape, and each accounted 
for at least 10% for the remaining variation among individuals. The population from Ash 
Warm Springs differed markedly (larger protarsus relative to meso- and metatarsi, and 
smaller pro-, meso-, and metatarsi relative to body length and synthlipsis) from the other 
populations, suggesting the need for systematic revision of its subspecific affiliation. In 
contrast, populations of A. m. heidemanni Montandon and A. m. minor La Rivers were 
similar to the other populations of the nominate subspecies, suggesting their subspecific 
recognition is questionable. The distinctiveness of the population from Ash Warm Springs 
as an isolated remnant of the historically connected pluvial White River intimates that it 
may be a taxon in the process of specific differentiation. 


Key Words: Naucoridae, Ambrysus mormon, morphometrics, phenetics, shape 


Ambrysus mormon Montandon is the Mexico. Throughout its range, A. mormon 
most widespread species of the genus inthe occurs in diverse habitats ranging from cold 
United States, ranging from Oregon and montane streams to thermal spring efflu- 
Idaho east to South Dakota, and south ents. Typically, it is found in slow water 
through Arizona and New Mexico into’ near margins of gravel-bottomed streams 


534 


(Usinger 1946). Four subspecies currently 
are recognized (La Rivers 1971): A. m. aus- 
tralis La Rivers in Mexico, southern Texas, 
and New Mexico; A. m. heidemanni Mon- 
tandon in thermal runoffs in Yellowstone 
National Park; A. m. minor La Rivers in a 
thermal spring in Idaho; and the nominate 
subspecies throughout the remainder of the 
range in the United States from Idaho and 
South Dakota south to Arizona and New 
Mexico. 

Because of their aquatic habitat require- 
ments, occurrence in otherwise arid land- 
scapes, and limited dispersal abilities, nau- 
corids in the southwestern United States are 
characterized by a high level of endemism. 
For example, A. relictus Polhemus and Pol- 
hemus and A. amargosus La Rivers, which 
is listed as a federally endangered species, 
occur only in several fragile streams in a 
western Nevada desert oasis; A. funebris La 
Rivers occurs only in Death Valley; and 
Limnocoris moapensis (La Rivers) occurs 
only in the vicinity of Moapa, Nevada. 

An isolated population of A. mormon at 
Ash Warm Springs, Nevada, possesses dis- 
crete morphological characteristics that dif- 
fer from those of other populations of the 
species. As a parallel study to an analysis 
of discrete characters (J. T. Polhemus, in 
litt.), we present an analysis of shape-relat- 
ed variation in A. mormon to determine the 
degree of divergence of the Ash Warm 
Springs population from other populations 
of the nominate subspecies, as well as from 
A. m. minor and A. m. heidemanni. 


MATERIALS AND METHODS 


A suite of 15 external mensural charac- 
ters (body length and width; head length 
and width; synthlipsis; pronotal length; 
lengths of pro-, meso-, and metathoracic fe- 
mur, tibia, and tarsus), previously deter- 
mined to be effective in discriminating 
among naucorid taxa (Sites and Willig 
1994a, b), was measured for adult speci- 
mens of 13 populations of A. mormon (Ta- 
ble 1, Fig. 1). Body length was measured 
from the tip of the labrum to tip of the ab- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


domen; body width, head length, head 
width, and all leg segments were longest 
distances; pronotal length was measured 
along the midline. Meso- and metanotal 
lengths were not included because it is dif- 
ficult to obtain accurate measurements 
without dissection of specimens. Each of 
these nota subducts below the preceding 
notum, and the visible length is variable 
and dependent on the degree of thoracic 
flexion. Generally, 10 specimens of each 
sex from each population were measured; 
however, for four populations, fewer than 
10 specimens were available (Eel River, 
Hot Creek Falls, Utah, and Yellowstone). 
All data were transformed to natural loga- 
rithms to evaluate more effectively the con- 
tribution of shape (see Sites and Willig 
1994a, b) to interpopulational differences. 
Statistical analyses were executed using 
SPSS (1990). Voucher specimens are de- 
posited in the Enns Entomology Museum, 
University of Missouri-Columbia, and the 
John T. Polhemus Collection. 

Data from labels of 18 of the 19 speci- 
mens of A. m. minor provided insufficient 
detail concerning site of collection (Table 
1, HCF) to ascribe individuals to the type 
locality for the subspecies with certainty. 
More specifically, a single specimen from 
the type locality of Hot Creek Falls is in- 
cluded in the analysis, along with a series 
of 18 specimens labeled “nr. Bruneau.” 
Therefore, the morphometric affinities of 
these specimens with that of the known to- 
potypic specimen were determined using 
discriminant function analysis with groups 
defined by combinations of sex and popu- 
lation. 

Two-way multivariate analysis of vari- 
ance (MANOVA) evaluated differences 
among populations and between sexes 
based on mensural characters. Univariate, 
two-way analyses of variance (ANOVAs) 
were performed to assess the contribution 
of each particular character to multivariate 
group differences. To minimize the likeli- 
hood of overestimating the significance of 
individual characters that compose a large 


VOLUME 102, NUMBER 3 


535 


Table 1. Acronyms and collection data for populations of Ambrysus mormon. Numbers of measured male 
and female specimens, respectively, appear parenthetically below each acronym. 


AWS 
(10,10) 


NEVADA: Lincoln Co. 
Ash Warm Springs 

3750 ft. elev.; 36°C 

21 July 1992; CL 2711 

J. T. and D. A. Polhemus 


COL 
(10,10) 


COLORADO: Pueblo Co. 
Burnt Mill Creek 
CL2683; 19-V-1992 

J. T. Polhemus 


EEL 
(8,10) 


CALIFORNIA: Mendocino Co. 
Eel River at Bell glen nr. Leggett 
62°F; 2-VII-1959 

R. K. Allen 


HCF 
(0,1) 


IDAHO: Owyhee Co. (1) 
Hot Cr. Falls 

9-TX-1965 

E. J. Allen 


HCF 
(10,8) 


IDAHO: Owhyee Co. (18) 
nr. Bruneau 

10-X-1975 

A. D. Allen 


IDA 
(10,10) 


IDAHO: Owyhee Co. 
Bruneau Riv. ca. 7 mi 
SSE Bruneau 

T7S ROE sect. 26 

22 Sept. 1988 

R. S. Zack, coll. 


INY 
(10,10) 


CALIFORNIA: Inyo Co. 
Shoshone Hot Spring 

CL 2904 

coll: J. T. and D. A. Polhemus 


MOA NEVADA: Clark Co. 
(10,10) Warm Springs; 32°C 

27 August 1989 

coll: J. A. Back 
NMX NEW MEXICO: Lincoln Co. 
(10,10) Rio Hondo 2 mi E Hondo 

23 Sept. 1988; 12°C 

coll: R. W. Sites 

veg. in flow at margin 
NYE NEVADA: Nye Co. 
(10,10) Hot Creek Spring 

7 mi. W. Hwy 93 

30°C; CL2898 

coll: J. T. and D. A. Polhemus 
SDA SOUTH DAKOTA: Fall River Co. 
(10,10) Hot Springs; 2 mi NW Hot Water 

22 June 1940; H. C. Severin 
SHA CALIFORNIA: Shasta Co. 
(10,10) Rt. 44; Cow Ck nr Palo Cedro 

28 Aug 1991; R. S. Zack and 

M. A. Valenti, collrs. 
UTA — UTAH: Washington Co. 
(10,8) North Creek nr. Virgin 

16-X-1982 

D. A. Polhemus 
YST WYOMING: Yellowstone Nat. Pk. 
(9,10) CL 556; 15-VIII-1972 


J. T. Polhemus 


suite of attributes (Holm 1979, Rice 1989), 
the Bonferroni sequential adjustment was 
applied to each morphometric character be- 
fore ascribing statistical significance to uni- 
variate analyses. Discriminant function 
analysis (DFA) simultaneously maximized 
intergroup differences and minimized intra- 
group variation among individuals by ad- 
justing the linear combination of variables 
in each of a number of orthogonal axes. 
Pairwise F-tests associated with DFA de- 
termined which populations differed signif- 
icantly from each of the other populations. 


The subsequent classification phase of DFA 
then assigned each specimen to a popula- 
tion based on the linear combination of var- 
iables from each discriminant function axis. 
Percent of correct assignments was used as 
a separate measure of morphometric dis- 
tinction among populations. Because DFA 
reconstitutes variables to maximize differ- 
ences among groups, natural relationships 
can become distorted in multidimensional 
space. In contrast, principal components 
analysis (PCA) was used as a data reduction 
technique to retain natural distance relations 


Janes AL 
in analyses. 


among individuals in multidimensional 
space and to view differences among pop- 
ulations with regard to shape and size for 
males and females separately. If the mag- 
nitude of differences in shape among pop- 
ulations is a consequence of the degree of 
isolation derived from geographic distance, 
then a correlation should exist between a 
matrix of pairwise linear distances between 
sites and a matrix of pairwise morphometric 
distances based on population centroids 
(PC2 through 4). We tested this hypothesis 
of differentiation by distance separately for 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Range (shaded) of Ambrysus mormon in the United States and localities (black circles) represented 


males and females using Mantel correlation 
analysis (Fortin and Gurevitch 1993, Manly 
1994, Sokal and Rohlf 1995). Via a sepa- 
rate Mantel analysis, we evaluated the de- 
gree to which interpopulational differences 
in shape between males and females were 
similar, regardless of geographic correlates. 


RESULTS 


The 18 specimens labeled “‘nr. Bruneau”’ 
of uncertain subspecific association likely 
represent A. m. minor because they formed 
a well-defined cluster in morphometric 


VOLUME 102, NUMBER 3 


Table 2. Significance levels (P)* of the character 
suite (MANOVA) and each character separately (AN- 
OVA) in distinguishing between sexes and among 13 
populations of Ambrysus mormon. 


Population Sex Pop X Sex 
MANOVA <0.001 <0.001 <0.001 
ANOVA 
Body length <0.001 <0.001 0.001 
Body width <0.001 <0.001 0.180 ns 
Head length <0.001 <0.001 0.908 ns 
Head width 0.001 <0.001 0.922 ns 
Synthlipsis <0.001 <0.001 0.121 ns 
Pronotum <0.001 <0.001 0.418 ns 
Profemur <0.001 <0.001 0.099 ns 
Protibia <0.001 <0.001 <0.001 
Protarsus <0.001 <0.001 0.003 
Mesofemur <0.001 <0.001 0.427 ns 
Mesotibia 0.001 <0.001 0.353 ns 
Mesotarsus <0.001 <0.001 0.099 ns 
Metafemur <0.001 <0.001 0.018 ns 
Metatibia <0.001 <0.001 0.009 ns 
Metatarsus <0.001 <0.001 0.437 ns 


4 Bonferroni sequential adjustments within a column 
for the 15 univariate characters corroborated signifi- 
cant (a = 0.05) group differences in all situations ex- 
cept those marked ns. 


space that was distinct from that of speci- 
mens representing the population of A. m. 
mormon in the Bruneau River (IDA). In ad- 
dition, each specimen was assigned to the 
correct population in the classification 
phase of DFA, further substantiating the 
morphometric distinction of these speci- 
mens. All 19 specimens match discrete 
characteristics listed for the subspecies by 
La Rivers (1963) and are small; this is typ- 
ical of naucorids that develop in high en- 
vironmental temperatures (Sites et al. 
1996). Differences among populations (Ta- 
ble 2) depended on sex (MANOVA, Pop- 
ulation by Sex interaction, P = 0.001). 
Three characters (lengths of body, protibia, 
and protarsus) contributed significantly (P 
<= 0.05) to the interaction (Table 2). All oth- 
er characters contributed significantly to 
differences (P = 0.001) among populations 
or between sexes in a consistent fashion. 
Pairwise F-tests revealed very highly 
significant morphometric differences (P = 


3)3)7/ 


0.001) between each possible pair of pop- 
ulations (78 pairwise contrasts per sex). 
The classification phase of DFA assigned 
all but two (98.43%) male specimens and 
all but one (99.21%) female specimen to 
the correct population. Clearly, intrapopu- 
lational morphometric variation was con- 
siderably less than interpopulational differ- 
ences. 

Principal components analysis effectively 
disassociated aspects of size (PC 1) and 
shape (other axes) from morphological var- 
iation among individuals. The loading co- 
efficient of each character was positive on 
PC 1 (Table 3) and accounted for 92.2% of 
the interindividual variation. Three addi- 
tional axes each accounted for at least 
10.0% of the shape-related variation. Of 
these shape axes, PC2, PC3, and PC4 ac- 
counted for 33.5%, 20.5%, and 10.4% of 
the shape-related variation among individ- 
uals, respectively. Males and females show 
correlated patterns of interpopulational var- 
iation in shape (Mantel analysis; P = 
0.015). This is reinforced by results from 
PCA in which the interpopulational disper- 
sion with respect to shape is similar in 
males and females (Fig. 2). The main dif- 
ference between the sexes is captured by 
PC2 (lengths of meso- and metatarsi rela- 
tive to protarsus): females from a particular 
population have larger scores on PC2 than 
do corresponding males. Nonetheless, the 
degree of differentiation in shape was un- 
related to geographic distance between pop- 
ulations for males (Mantel analysis; P = 
0.604) and females (Mantel analysis; P = 
0.278). Factors other than distance, per se, 
must be affecting interpopulational varia- 
tion in shape in this species. 

Regardless of sex, the attribute of shape 
that distinguishes the Ash Warm Springs 
population from the other populations of A. 
mormon ‘is its relatively elongate protarsi 
compared to meso- or metatarsi (PC2). To 
a lesser degree, this same attribute of shape 
distinguished populations of MOA and 
NYE from the remainder of populations, es- 
pecially when considering females. Addi- 


538 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 

Table 3. Principal component loadings (correlations) for axes representing size (PC1) and appreciable attri- 
butes of shape (PC2 through 4). 

Character PCl PeZ PC3 PC4 

Body length 0.964 —0.031 —0.143 —0.168 
Body width 0.986 —0.006 —0.057 Oss 
Head length 0.978 —0.019 —0.074 —0.024 
Head width 0.977 —0.006 —0.058 —0.006 
Synthlipsis 0.925 0.164 —0.246 0.109 
Pronotum length 0.964 —0.098 —0.108 —0.006 
Profemur length O97S —0.054 SOOT 0.175 
Protibia length 0.970 —0.099 0.014 0.174 
Protarsus length 0.883 —0338 0.258 —0.002 
Mesofemur length 0.992 —0.014 —0.007 0.002 
Mesotibia length 0.986 —0.068 0.046 —0.006 
Mesotarsus length 0.880 0.398 0.203 0.020 
Metafemur length 0.992 —0.016 0.009 —0.044 
Metatibia length 0.985 —0.037 0.065 —0.063 
Metatarsus length 0.936 0.250 0.144 —0.040 
Eigenvalue 13.826 0.393 0.240 0.122 

% variance (total) 0.922 0.026 0.016 0.008 

% variance (shape) —_ 0.335 0.205 0.104 


tional shape variation that distinguishes all 
populations (PC3) is related to length of tar- 
si relative to body size and synthlipsis. In 
particular, the relative lengths of tarsi in the 
three populations from White River (AWS, 
MOA, and NYE) are smaller than those in 
the other populations. 


DISCUSSION 


Recently, Sites and Willig (1994a, b) 
demonstrated that the above character suite 
was effective at discriminating among gen- 
era and species of naucorids. This character 
suite is not only diacritical in distinguishing 
among taxa but discriminates among con- 
specific populations of A. mormon. None- 
theless, the particular components of shape 
that distinguished 22 species of Ambrysus 
(Sites and Willig 1994a) are different than 
those that distinguished populations of A. 
mormon. The positions of populations of A. 
mormon in multidimensional morphometric 
space do not reflect currently established 
subspecific designations within the species. 
More specifically, A. m. heidemanni (YST) 
and A. m. minor (HCF) occur in close mor- 
phometric proximity to members of the 
nominate subspecies (all other populations), 


and occur within the range of shape vari- 
ability encompassed by the nominate sub- 
species (Fig. 2). Moreover, populations of 
the nominate subspecies from Ash Warm 
Springs, Moapa, and New Mexico (AWS, 
MOA, NMX) are considerably more diver- 
gent based on shape-related variation (Fig. 
2) than either of the two other currently rec- 
ognized subspecies included in this analysis 
(A. m. heidemanni, YST; A. m. minor, 
HCF). Further, discrete characters defining 
A. m. heidemanni, which occurs in the 
warm waters of the Yellowstone geyser ba- 
sins, may be expressed in other warm water 
populations of nominate A. mormon, which 
led La Rivers (1951) to question the sub- 
specific affiliation of this population. Be- 
cause shape-related variation among popu- 
lations does not reflect the currently ac- 
cepted intraspecific taxonomy of A. mor- 
mon, and discrete characters offer only 
equivocal support, re-evaluation of subspe- 
cific status is warranted for those popula- 
tions. 

Populations from Utah, Yellowstone, 
Colorado, and South Dakota; as well as 
those from Idaho and New Mexico (Sites 
et al. 1996), reflect a wide range of envi- 


VOLUME 102, NUMBER 3 539 


Males 


UTA 


SHA 


— 


o PC3 


2 
UTA 
1 SHA 
EEL 
re SDA 
(9 NmMx© OL 
jae 
H 
[ICE vy IDA 
NYE 
-1 MOA 
~ BS Sheauacnba ce 2 1 0 1 2 
<9 22 aS Sashes & 


PC2 


Fig. 2. Plots of 13 population centroids, derived from a principal components analysis of 15 mensural 
characters, illustrate intraspecific size (PC1) and shape (PC 2 and 3) variation for male and for female A. mormon. 
Eleven populations represent nominate A. mormon, whereas HCF and YST populations (in boxes) represent A. 
m. minor and A. m. heidemanni, respectively. 


540 


ronmental temperatures during develop- 
ment, yet the positions of group centroids 
(Fig. 2) indicate that these populations 
share similar attributes of shape. Although 
thermal environment does influence allo- 
metric relationships during ontogeny (Sites 
et al. 1996), the greatest degree of shape- 
related variation in A. mormon has been 
achieved independently of thermal environ- 
ment during ontogeny. 

The population of A. m. minor (HCF) ex- 
hibits morphometric attributes distinct from 
those of the nearby population of nominate 
A. mormon in the Bruneau River (IDA), 
based on a pairwise F-test from DFA. De- 
spite the close proximity of Hot Creek Falls 
to Bruneau River (~1 km), distinct mor- 
phologies characterize the populations of 
the two sites. A series of unnamed hot 
springs occurs along the Bruneau River, and 
A. mormon may occur in them. Because of 
local irrigation demands, the water table has 
declined recently, resulting in a concomi- 
tant reduction in the size of the springs. 
During the last known visit to these springs 
(1991), the pool at the base of Hot Creek 
Falls, known as Indian Bathtub, was re- 
duced tremendously in size (~1.0 m diam 
x 0.3 m depth) and persisted only because 
a piece of plastic had been placed on the 
downstream side of the pool, probably to 
allow bathing; the actual falls no longer ex- 
ists (R. S. Zack, personal communication). 
Continued reduction in the water table may 
threaten A. m. minor to the point that it may 
soon become extinct. 

The White River system in eastern Ne- 
vada extends from the White Pine Moun- 
tains near Ely south to the Moapa River 
near Lake Mead. The lower end of the 
White River near Moapa is known as the 
Muddy River. The White River system is of 
hydrographic and faunistic interest because 
of its recently restricted watercourse and 
isolated fish populations. In the Pliocene 
and at least the early Pleistocene epochs, 
the pluvial White River was continuous 
throughout its length to an isolated lake, 
now represented by the Moapa River (Gil- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


bert 1893, Carpenter 1915), which persists 
as a tributary of the Colorado River. Since 
that time, extensive desertification of the 
Great Basin has occurred (King 1958), sig- 
nificantly reducing the extent of the White 
River. As a result, ~200 miles of dry river 
bed now cross the parched desert, isolating 
the Recent White River (north) from the 
Moapa River (south). Despite drastic reduc- 
tions in surface water, sporadically distrib- 
uted springs and effluent fragments of the 
pluvial White River persist. 

The fish fauna of the White River is dis- 
tinct and includes an endemic cyprinid ge- 
nus, an endemic species of Crenichthys, 
and several indigenous species and subspe- 
cies (Hubbs and Miller 1948). Subspecific 
affinities of isolated fish populations in 
these springs and effluents strongly suggest 
that these habitats were continuous in the 
past. Ambrysus mormon also occurs in 
many of these aquatic fragments, a result of 
its almost certain occurrence in the pluvial 
White River and subsequent isolation fol- 
lowing landscape wide desertification. 
Three populations included in our morpho- 
metric assessment represent White River 
isolates [AWS, MOA, and NYE (see Table 
1)]. Specimens of the Ash Warm Springs 
population exhibit structural features that 
are divergent from “typical”? A. mormon, 
including reduction in posterolateral con- 
nexival spines, narrow embolium, and dif- 
ferences in genitalic features. 

Evaluation of discrete-state characters of 
the isolated Ash Warm Springs population, 
combined with the attributes of shape of 
other members of the species, suggests that 
this population may warrant taxonomic rec- 
ognition, at least at the subspecific level. 
Although the length of time that it has been 
separated from the parent population from 
White River is uncertain, clear differences 
in phenotype, including shape, have accu- 
mulated in isolation. Even if speciation is 
not yet complete, the population likely will 
continue to diverge from the other popula- 
tions of A. mormon because of the effects 
of drift and natural selection in environ- 


VOLUME 102, NUMBER 3 


mental isolation. If divergence has not pro- 
gressed to the point at which reproductive 
incompatibility exists between the Ash 
Warm Springs population and the remain- 
der of A. mormon, then we are witnessing 
speciation in action in this remnant popu- 
lation of A. mormon. Although the Nye Co. 
and Moapa populations also represent iso- 
lates along the White River system, these 
populations have not diverged from typical 
morphometric form of A. mormon as radi- 
cally as has the population at Ash Warm 
Springs (Fig. 2). Further studies involving 
discrete state characters are needed to re- 
solve the intraspecific taxonomic status of 
these populations of A. mormon. 


ACKNOWLEDGMENTS 


We thank John T. Polhemus (Englewood, 
CO) and Richard S. Zack (Washington 
State University) for the loan and gift of 
selected specimens for this project, and Ri- 
chard S. Zack for details concerning local- 
ities of A. m. minor. We also thank J. E. 
McPherson (Southern Illinois University) 
and John T. Polhemus for critical reviews 
of this manuscript. The Enns Entomology 
Museum provided optical and computing 
facilities. This research was completed 
while MRW was a Sabbatical Fellow at the 
National Center for Ecological Analysis 
and Synthesis, a center funded by NSF 
(DEB-9421535), the University of Califor- 
nia at Santa Barbara, and the state of Cal- 
ifornia. A Developmental Leave was pro- 
vided to MRW by the Provost’s Office, 
Texas Tech University. Funding for RWS 
was provided in part by University of Mis- 
souri project #PSSLO232. This is Missouri 
Agricultural Experiment Station Journal Se- 
ries paper 12,911. 


LITERATURE CITED 


Carpenter, E. 1915. Ground water in southeastern Ne- 
vada. United States Geological Survey, Water- 
Supply Paper 333: 1—90, figs. 1-9, pls. 1-2. 

Fortin, M.-J. and J. Gurevitch. 1993. Spatial statistics: 
Analysis of field experiments, pp. 319-341. In 
Scheiner, S. M. and G. Gurevitch, eds., Design 


541 


and Analysis of Ecological Experiments. Chap- 
man and Hall, New York. 445 pp. 

Gilbert, C. H. 1893. Report on the fishes of the Death 
Valley Expedition collected in southern California 
and Nevada in 1891, with descriptions of new spe- 
cies. North American Fauna 7: 229-234, pls. 5—6. 

Holm, S. 1979. A simple sequentially rejective multi- 
ple test procedure. Scandinavian Journal of Statis- 
tics 6: 65-70. 

Hubbs, C. L. and R. R. Miller. 1948. The Great Basin, 
with emphasis on glacial and postglacial times. II. 
The zoological evidence: Correlation between fish 
distribution and hydrographic history in the desert 
basins of western United States. Bulletin of the 
University of Utah 38(20): 18-166 + 1 map. 

King, P. B. 1958. Evolution of modern surface features 
of western North America. pp. 3—60. Jn Hubbs, 
C. L., ed. Zoogeography. American Association 
for the Advancement of Science Publication Num- 
ber 51, 509 pp. 

La Rivers, I. 1951. A revision of the genus Ambrysus 
in the United States (Hemiptera: Naucoridae). 
University of California Publications in Entomol- 
ogy 8: 277-338. 

. 1963. Two new Ambrysi (Hemiptera: Nau- 

coridae). Biological Society of Nevada Occasional 

Papers 1: 1-7. 

. 1971. Studies of Naucoridae (Hemiptera). Bi- 
ological Society of Nevada Memoirs 2, ii + 120 
pp- 

Manly, B. FE J. 1994. Multivariate statistical methods: 
A primer, 2nd ed. Chapman and Hall, New York. 
159 pp. 

Rice, W. R. 1989. Analyzing tables of statistical tests. 
Evolution 43: 223-225. 

Sites, R. W. and M. R. Willig. 1994a. Interspecific af- 
finities in Ambrysus (Hemiptera: Naucoridae). 
Proceedings of the Entomological Society of 
Washington 96: 527-532. 

. 1994b. Efficacy of mensural characters in dis- 
criminating among species of Naucoridae (Insec- 
ta: Hemiptera): Multivariate approaches with an 
ontogenetic perspective. Annals of the Entomo- 
logical Society of America 87(6): 803-814. 

Sites, R. W., M. R. Willig, and R. S. Zack. 1996. Mor- 
phology, ontogeny, and adaptation of Ambrysus 
mormon (Hemiptera: Naucoridae): Quantitative 
comparisons among populations in different ther- 
mal environments. Annals of the Entomological 
Society of America 89: 12-19. 

Sokal, R. R. and FE J. Rohlf. 1995. Biometry: The Prin- . 
ciples and Practice of Statistics in Biological Re- 
search, 3rd ed. W. H. Freeman, New York. 887 pp. 

SPSS, Inc. 1990. The SPSS Base System User’s Guide. 
SPSS, Inc., Chicago. 520 pp. 

Usinger, R. L. 1946. Notes and descriptions of Ambry- 
sus Stal with an account of the life history of Am- 
brysus mormon Montd. (Hemiptera, Naucoridae). 
University of Kansas Science Bulletin 31: 185-210. 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 542-548 


NOTES ON NEOTROPICAL SPECIES OF TETHINA HALIDAY 
(DIPTERA: TETHINIDAE) 


GEORGE A. FOSTER AND WAYNE N. MATHIS 


Department of Entomology, National Museum of Natural History, Smithsonian Insti- 
tution, Washington, D.C. 20560-0169, U.S.A. (e-mail: agromyza@msn.com and 
mathis. wayne @nmnh.si.edu) 


Abstract.—Neotropical beach-fly species (Diptera: Tethinidae) not included in a pre- 
vious faunal revision of the Caribbean, Gulf of Mexico, and Bermuda are reviewed, 
including description of the following new species (type locality in parenthesis): Tethina 
robusta (Chile. Osorno: Puchatrihue). Two new synonyms are also proposed (junior syn- 
onyms cited first): Rhicnoessa sonorensis Melander = Tethina albula Loew and Tethina 
setulosa Malloch = Tethina spinulosa Cole. A revised key to the Neotropical species of 


Tethina and a diagnosis for the senior synonyms are provided. 


Key Words: 


Since revising the Tethinidae from the 
Caribbean, Gulf of Mexico, and Bermuda 
(Foster and Mathis 1998), we have studied 
additional Neotropical specimens that have 
revealed a striking new species from Chile 
and two synonyms of widespread species. 
This paper reports these discoveries. Here- 
in, we present a revised key to the Neo- 
tropical species of Tethina and diagnoses 
for the species for which synonyms have 
been discovered. This paper is written with- 
in the context of our recent revision (Foster 
and Mathis 1998), and further details are 
found in that reference. 

The discovery of the two new synonyms 
reported herein re-emphasizes our previous 
observation (Foster and Mathis 1998) that 
most coastal marine species of Tethinidae 
have widespread distributions. Although we 
studied most species of Tethina that were 
known to occur in the New World for our 
Caribbean study, we did not examine two 
species, T. spinulosa Cole and T. sonorensis 
(Melander), as both were reported to occur 
only in western North America and our 


Diptera, Tethinidae, Tethina, neotropics 


study concerned the Caribbean and adjacent 
areas. Our study of these two species re- 
veals that they too are conspecific with oth- 
er, widespread species as we document be- 
low. 

Methods.—The descriptive terminology, 
with the exceptions noted in Mathis and 
Munari (1996), follows that of McAlpine 
(1981). We have followed the terminology 
for most structures of the male terminalia 
that other workers in Tethinidae have used 
(see references in Mathis and Munari 
1996). The terminology for structures of the 
male terminalia is labeled on Figs. 2—4. The 
description of the new species is based pri- 
marily on its holotype. 

States of Mexico are abbreviated as fol- 
lows: Baja California Norte (BCN), Quin- 
tana Roo (QNR), Sonora (SON), and Ta- 
basco (TAB). 


KEY TO NEOTROPICAL SPECIES OF TETHINA 


1. Gena high, at least 0.5 eye height 
— Gena short, less than 0.5 eye height ...... 2; 
Apex of scutellum with yellowish to reddish 


N 


VOLUME 102, NUMBER 3 


spot (may vary in size but always obvious) 
T. xanthopoda (Williston) 
— Apex of scutellum uniformly gray microto- 
ITICTILOSE senate PeR NE ot oe eee een i en rade aca 3 
3. Mid- and hindtibiae black on apical ; first 3 
tarsomeres white to pale yellow ......... 
bn melee Sh ye A. T. albitarsa Foster and Mathis 
— Mid- and hindtibiae yellow; first 3 tarsomeres 
VEUOWA: crops cok tuene 2 ouseebnes sataa apes 4 
4. Hindfemur of d not particularly swollen... 5 
Hindfemur of d distinctly swollen 
5. Center of gena with elongate, shiny area ap- 
parently free of microtomentum (fig. 24 in 
Foster and Mathis 1998); surstylus armed 
with many short, well-developed, thick tooth- 
hike ‘setulae 3b 3.-54 5; T. texana (Melander) 
— Gena uniformly microtomentose (d  un- 
known) T. insulans Curran 
6. Surstylus curved anteriorly, ending in an 
acute point, densely setulose with well-devel- 
oped setae over most of surface 


T. cohiba Foster and Mathis 
—  Surstylus paddle shaped, with well-developed 

setae along margin only .... 7. spinulosa Cole 
7. Body distinctly gray to grayish brown .... 8 

Bodyawihitishtes £e;-ttet eae ses kt.) fee 9 
Ss )libiaexyellow) 3. =~ = - T. willistoni (Melander) 
bide yOray. smi CrOLOmentOSeCmesyes “fate a oe 
T. robusta, new species 
9. Surstylus in lateral view curved anteriorly .... 
T. albula Loew 
—  Surstylus in lateral view straight........ 10 
10. Surstylus in posterior view with median mar- 

gin curved and rounded, sparsely setulose ... . 
T. lisae Foster and Mathis 
—  Surstylus in posterior view with median mar- 

gin straight, bearing dense row of setulae .... 
T. bermudaensis (Melander) 


Tethina insulans Curran 


Tethina insulans Curran 1932: 358.—Fos- 
ter 1976b: 2 [Neotropical catalog].— 
Mathis and Munari 1996: 17 [world cat- 
alog]. 


Diagnosis.—Body length 3 mm, gener- 
ally gray, microtomentose; setae and setulae 
black. Gena moderately high, slightly less 
than 0.5 eye height. Presutural acrostichal 
setae 4; scutellum uniformly gray, lacking 
a dorsal spot. Femora gray, microtomento- 
se; hindfemur of male similar to or only 
slightly more swollen than fore- and mid- 
femora; tibiae yellow. Male terminalia un- 
known. 


543 


Distribution—WNeotropical: Ecuador (Ga- 
lapagos Islands). 

Remarks.—We studied the apparent al- 
lotype female and only known specimen of 
T. insulans, which is deposited in the 
AMNH. The holotype male was originally 
deposited in the Zoological Museum, Uni- 
versity of Oslo, Oslo, Norway [type local- 
ity: Ecuador. Galapagos Islands: Floreana, 
Post Office Bay (seaside)], and may now be 
lost. 

Curran’s original description character- 
izes the allotype well and is not repeated 
here. The allotype, which is covered with 
white particles, is in poor condition (head 
partially collapsed, glue covering mouth- 
parts, thorax very greasy, and abdomen is 
wrinkled and partially collapsed), making it 
difficult to study. 

This species is apparently related to T. 
milichioides, as Curran noted in the original 
description (Curran 1932:359): “‘[This spe- 
cies] In Melander’s key to Rhicnoessa 
(Tethina), traces to milichiodes [sic] Melan- 
der but differs in having reddish antennae, 
etc.”” Tethina milichioides is known from 
western United States (Mathis and Munari 
1996). Although the setae are not as short 
as in T. milichioides, this species shares 
many of characters of the milichioides 
group (Foster 1976a). We are satisfied that 
T. insulans is not conspecific with any 
known Neotropical species. 


Tethina albula (Loew) 
(Figs. 8-10, 13 in Foster and Mathis 
1998) 


Rhicnoessa albula Loew 1869: 44.—Mal- 
loch 1913: 147 [citation].—Melander 
1913: 298 [key].—Hendel 1934:43 [key], 
46 [citation].—Hennig 1937: 140 [cita- 
tion].—Melander 1952: 202 [citation]. 

Tethina albula: Curran 1934: 330 [generic 
combination].—Vockeroth 1965: 727 
[Nearctic catalog].—Prado and Tavares 
1966: 431 [revision].—Foster 1976b: 2 
[Neotropical catalog]—Mathis and Mu- 
nari 1996: 14 [world catalog].—Foster 
and Mathis 1998: 609—611 [revision]. 


544 


Rhicnoessa sonorensis Melander 1952: 
207.—Cole 1969: 387 [distribution, di- 
agnosis].—Foster 1976b: 2 [lectotype 
designation]. New synonym. 

Tethina sonorensis: Foster 1976b: 2 [ge- 
neric combination, Neotropical catalog]. 


Diagnosis.—This species is distin- 
guished from congeners by the following 
combination of characters: Body length 
1.60—3.15 mm; body with gray to whitish 
gray microtomentum; setae generally 
white to mostly black (Guyana specimens); 
gena high, greater than 0.5 eye height; 4 
irregular rows of acrostichal setulae; scu- 
tellum uniformly gray, lacking a spot; fem- 
ora mostly yellow to mostly gray; hindfe- 
mora of male similar to or only slightly 
more swollen than fore- and midfemora; 
tibiae yellow; basal 4 tarsomeres yellow, 
apical tarsomere brown; surstylus articu- 
lated with and broadly attached to epan- 
drium, narrowly spatulate in posterior view 
(Figs. 8, 10 in Foster and Mathis 1998), 
length 2.5 width, apex broadly rounded; 
median margin bearing irregular row of 
sparse setulae along entire length, setulae 
moderately well developed; surstylus in 
lateral view (Figs. 9-10 in Foster and 
Mathis 1998) narrow, height 3.5 width, 
gently curved anteriorly, basal portion pro- 
duced anteriorly as a lateral lobe that bears 
a patch of setulae mesally; aedeagus thick, 
straplike, bearing dense velvety hairlike 
pubescence on dorsal surface. 

Type material.—The syntype series, la- 
beled “‘Loew Collection’? and comprising 
one male (only two legs and a left wing 
remain) and four females (one bearing a 
red “‘Type”’ label (13444); MCZ), does not 
allow for accurate and reliable identifica- 
tion of this species. Osten Sacken, how- 
ever, collected and retained a male (head 
missing) from the type locality (Newport, 
Rhode Island) when he collected the type 
series. Osten Sacken’s practice was to re- 
tain a few specimens of species represent- 
ed by a long series, sending the majority 
to Loew for description. That retained, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


headless male, which is presumably con- 
specific with the type series, was identi- 
fied, dissected, and is the basis for our di- 
agnosis of this species. 

The lectotype male of Rhicnoessa son- 
orensis (designated by Foster, 1976b:2) is 
labeled ‘“‘SonoraMEXICO RockyP[oin]t 
Marsh 21 April ’47 [1947] A.L. Melander/ 
ALMelander Collection 1961 [stippled 
green band on right side of leabel]/Lecto- 
type Tethina sonorensis (Melander) George 
A. RFoster 1974 [handwritten; black sub- 
marginal border].’’ The lectotype is double 
mounted (minuten in a rectangular card), 
is in excellent condition (abdomen re- 
moved and dissected, the structures in an 
attached microvial), and is deposited in the 
USNM. 

Other specimens examined.—MEXICO. 
Sonora: Pt. Penasco, 21 Apr 1948, A. L. 
Melander (1 3, 3 2; USNM); Rocky Point 
Marsh, 21 Apr 1947, A. L. Melander (6 6, 
10 2; USNM). 

Distribution.—Nearctic: USA (DE, FL, 
MA, MD, NC, NY, SC, RI, VA). Neotrop- 
ical: Bahamas, Belize, Guyana, Mexico 
(QNR, SON), Trinidad, Turks and Caicos, 
West Indies (Grand Cayman). 

Remarks.—This is the first record of T. 
albula from the West Coast of the New 
World; we anticipate that it will be found 
to be more widespread. 


Tethina spinulosa Cole 
(Figs. 26, 28-29 in Foster and Mathis 
1998) 


Tethina spinulosa Cole 1923: 478.—Hen- 
del 1934: 41 [citation].—Vockeroth 
1965: 728 [Nearctic catalog].—Foster 
1976b: 2 [Neotropical catalog]_—Mathis 
and Munari 1996: 18 [world catalog]. 

Rhicnoessa spinulosa: Melander 1952: 202, 
208 [key, generic combination, citation]. 

Tethina setulosa Malloch 1934: 454.—Fos- 
ter 1976b: 2 [Neotropical catalog].— 
Mathis and Munari 1996: 18 [world cat- 
alog].—Foster and Mathis 1998: 624 [re- 
vision]. New synonym. 


VOLUME 102, NUMBER 3 
Rhicnoessa_ setulosa: Hennig 1937: 139 
[generic combination, citation]. 


Diagnosis.—This species is distinguished 
from congeners by the following combina- 
tion of characters: Body length 1.85—2.70 
mm; body generally with gray microtomen- 
tum; setae generally black; gena short, less 
than 0.5 eye height; 4 somewhat irregular 
rows of acrostichal setulae; scutellum uni- 
formly gray, lacking apical spot; femora 
mostly gray; hindfemora of male distinctly 
swollen, distinctly larger than fore- and 
midfemora; tibiae and basal 4 tarsomeres 
yellow, apical tarsomere brown; surstylus 
articulated with and broadly attached to 
epandrium, broadly spatulate in posterior 
view (Fig. 29 in Foster and Mathis 1998), 
length about equal to width, median margin 
bearing dense patch of robust setulae along 
entire length (Fig. 29 in Foster and Mathis 
1998); surstylus in lateroblique view (Fig. 
28 in Foster and Mathis 1998) broadly 
rounded, constricted anteriorly, external 
surface bearing numerous setulae; aedeagus 
thin, ribbonlike. 

Type material.—The holotype male of 
Tethina spinulosa is labeled ‘“[MEXICO. 
Baja California Norte:] Las Animas Bay[,] 
Gulf Cal. May 8 1921/EPVan Duzee Col- 
lector/HOLOTYPE spinulosa/ALLOTYPE 
spinulosa/Tethina spinulosa Type and allo- 
type [two specimens on separate points; 
type data taken from Arnaud 1979:345].” 
The holotype and allotype are double 
mounted (glued to separate paper points on 
same pin) and are deposited in the CAS 
(1356). We examined 30 paratypes, includ- 
ing 18 from the type locality, as follows: 
Mexico. Baja California Norte: Las Animas 
Bay, 8 May 1921, E. P- Van Duzee (4 ¢, 
14 2; USNM); Loreto, 19 May 1921, E. P. 
Van Duzee (8 3, 4 2; USNM). 

The holotype male of Tethina setulosa is 
labeled *“‘Angol [crossed out] Chile DSBul- 
lock/Tocopilla [Antofagasta] Ap. 10, [19]31 
Sea Beach [handwritten]/Type No. 50448 
U.S.N.M. [red; “50448” handwritten ]/Teth- 
ina setulosa Type Det. JRMALLOCH [spe- 


545 


cies name and “‘Type”’ handwritten; black 
submargin].’’ The holotype is directly 
pinned, is in good condition (abdomen re- 
moved and dissected, the parts are in an 
attached microvial), and is deposited in the 
USNM (50448). 

Other specimens examined.—ECUA- 
DOR. Galapagos: Isla Santa Cruz: Acade- 
my Bay, Darwin Research Station (beach 
and coastal rocks), 24 Jan 1964, D. Q. Cav- 
agnaro, R. O. Schuster (7 ¢, 6 2; USNM). 

MEXICO. Sonora: Guaymas (40 mi N), 
24 Nov 1951, J. J. Teas (3 6,4 9; USNM); 
Pto. de Lobos, 18-19 Mar 1974, W. Brown, 
V; Roth*S' 6; 3292 USNM): “Marsh: 21 
Apr 947, A. L. Melander (1 6; USNM). 

UNITED STATES. California. Orange: 
Balboa, 13 Jul 1940 (2 6; USNM); Corona 
del Mar, 25 Jul—-19 Nov, 1942-1949, A. L. 
Melander (5 36, 9 2; USNM); Doheny 
Park, 12 Oct 1951, A. L. Melander (3 6, 7 
2; USNM); Laguna Beach, 18 Jul-12-20 
Oct 1943, 1951, A. L. Melander (15 6, 9 
2; USNM); San Clemente, 13 Oct 1950, A. 
L. Melander (3 36, 4 2; USNM); Seal 
Beach, 26 Jul 1942, A. L. Melander (2 6, 
1 2; USNM). Los Angeles: Palos Verdes, 
15 Oct 1944, A. L. Melander (4 6,7 2; 
USNM). San Diego: Scripps, 8 Oct 1972, 
L. Chang (4 6, 2 2; USNM). 

Clipperton Island (10°17'N, 109°13'W; a 
French possession off the Pacific Coast of 
Mexico), 22 Aug 1958, C. E Harbison (15 
36,6 2; USNM). 

Distribution.—Widespread, primarily on 
the west coast of the New World. Nearctic: 
United States (CA). Neotropical: Chile 
(Tarapaea to Antofagasta), Clipperton Is- 
land, Ecuador (Galapagos), Mexico (BCN, 
SON, TAB). 

Remarks.—Although this species was 
known previously from Chile (Malloch 
1934) and later from the Caribbean coast of 
Mexico (TAB; Foster and Mathis 1998), we 
have identified numerous other specimens 
from the West Coast of the New World as 
noted above. The specimens from Clipper- 
ton Island, a French possession in the north 


546 


Pacific, approximately 1,100 km from the 
west coast of Mexico, are of note. 


Tethina robusta Foster and Mathis, 
new species 
(Figs. 1—4) 


Diagnosis.—This species is distinguished 
from congeners by the following combina- 
tion of characters: Body with gray micro- 
tomentum, thorax entirely gray; setae black; 
gena high, 0.62—0.75 eye height; 4 irregular 
rows of acrostichal setulae; scutellum uni- 
formly gray, lacking yellowish to reddish 
spot; femora distinctly gray; fore- and hind- 
femora of male distinctly swollen, distinctly 
larger than midfemora, with hindfemora 
distinctly larger than forefemora; tibiae 
black; basal 3 tarsomeres yellow, appearing 
velvety ventrally on fore- and hindleg; sur- 
stylus broadly spatulate and broadly at- 
tached to epandrium posteriorly, bearing 2 
rows of short setae along medial margin 
and scattered setae on posterior surface; 
epandrium with a large, triangular, ventral 
lobe nearly as long as surstylus, sparsely 
setulose on medial surface, bearing a row 
of short setae along posterior margin; ae- 
deagus very thin, ribbonlike. 

Description.—Body length 2.65—2.85 
mm; body with gray microtomentum; setae 
black. 

Head (Fig. 1): Setae black. Vertex gray, 
microtomentose; ocellar tubercle bearing 2 
ocellar setae and 1 shorter setula directly in 
middle; postocellar seta well developed; 
frons yellowish medially, grayish laterally, 
microtomentose; interfrontal setae 3, ante- 
rior pair proclinate, posterior 2 setae con- 
vergent; fronto-orbital setae as 3 conver- 
gent, proclinate inner setae and 4 divergent, 
outer setae; bearing 2 additional pairs of 
dorsally directed setae near base of anten- 
nae. Antenna dark brown except basal % of 
Ist flagellomere reddish brown; arista dark 
brown, sparsely pubescent. Face white mi- 
crotomentose; facial tubercle yellow, dor- 
sad of anteriormost seta; peristomal setae 
bearing 4 dorsally directed setae, 5th di- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


rectly anteriorly. Gena high, 0.62—0.75 eye 
height, white, microtomentose; postgena 
gray bearing pale setulae. Palpus yellow; la- 
bellum long, brown. 

Thorax: Entirely gray microtomentose; 
scutellum uniformly gray, lacking yellow- 
ish to reddish spot. Acrostichal setulae in 4 
irregular rows; dorsocentral setae 5 (2+3); 
proepisternum and proepimeron each bear- 
ing 1 seta. Wing with veins brown to yel- 
low except center of crossveins distinctly 
whitish; costa continuing strongly to vein 
M; vein R,,, and R,,,; divergent; veins R,,; 
and M parallel apically. Midtibia ventrally 
bearing several strong black setae in addi- 
tion to long apical seta; hindtibia bearing 1 
well-developed seta in addition to apical 
seta; coxae gray to brownish gray, bearing 
mostly pale setulae; femora entirely gray 
microtomentose, swollen, especially hind- 
femur, which is distinctly more swollen 
than fore- and midfemora; tibiae gray mi- 
crotomentose; basal 3 tarsomeres yellow, 
apical tarsomeres black. 

Abdomen: Setae and setulae black; distal 
margins of segments yellow, remainder 
concolorous with thorax. Male terminalia 
(Figs. 2-4): Surstylus broadly spatulate and 
broadly attached to epandrium posteriorly, 
bearing 2 rows of short setae along medial 
margin and scattered setae on posterior sur- 
face; epandrium with a large, triangular, 
ventral lobe nearly as long as _ surstylus, 
sparsely setulose on medial surface, bearing 
a row of short setae along posterior margin; 
aedeagus very thin, ribbonlike. 

Type material.—The holotype d is la- 
beled ‘“‘CHILE: Osorno Pr[ovince]: Puca- 
trihue, 27-30 January 1978 WNMathis/ 
HOLOTYPE Tethina robusta 6 Foster & 
W.N.Mathis USNM [red; species name and 
‘“*$ & Foster’ handwritten].’’ The holotype 
is double mounted (minuten in a block of 
plastic), is in excellent condition (a few se- 
tae with apices broken), and is deposited in 
the USNM. Three paratypes, all males, bear 
the same locality label data as the holotype. 

Distribution.—Neotropical: Chile. Osor- 
no: Pucatrihue. 


VOLUME 102, NUMBER 3 547 


a= hypandrium 


, 

Sa th 

Mat 

7 Pag 
‘ raat 
i can] 
1 x OE 
wk 
fetes 


Sebel ges -— hypandrial apodeme 


—— aedeagal apodeme 


SN \ 3 
“SS ventral lobe A 
surstylus 


Figs. 1-4. Tethina robusta: 1, Head, lateral view. 2, External male terminalia, posterior view (Chile. Orsono: 
Puchatrihue). 3, Same, lateral view. 4, Internal structures of male terminalia (aedeagus excluded), ventral view. 


548 


ACKNOWLEDGMENTS 


We express our sincere thanks to Dr. Da- 
vid A. Grimaldi (American Museum of 
Natural History, New York, New York) for 
loaning specimens in his care. Without his 
cooperation this study could not have been 
completed. 

Field work in Chile was greatly expedit- 
ed through the able and pleasant assistance 
of Oliver S. Flint, Jr. and Paul J. Spangler, 
and we are also grateful to David Challinor 
(former Assistant Secretary for Research, 
Smithsonian Institution) for financial sup- 
port to conduct this field work. Field work 
on St. Vincent, St. Lucia, Dominica was 
supported by a grant from the Research Op- 
portunity Fund, then administered by Stan- 
wyn G. Shetler (former Deputy Director of 
the National Museum of Natural History). 

The illustrations were carefully inked by 
Young T. Sohn. For reviewing a draft of this 
paper we thank Lorenzo Munari and Ray- 
mond J. Gagné. 


LITERATURE CITED 


Arnaud, P. H., Jr. 1979. A catalog of the types of Dip- 
tera in the collection of the California Academy 
of Sciences. Myia 1: vi+505 pp. 

Cole, F R. 1923. Expedition of the California Acade- 
my of Sciences to the Gulf of California in 1921. 
Diptera from the islands and adjacent shores of 
the Gulf of California. Il. General Report. Pro- 
ceedings of the California Academy of Sciences 
12(25): 457-481. 

Cole, E R. (with the collaboration of E. T. Schlinger). 
1969. The Flies of Western North America. Uni- 
versity of California Press, Berkeley and Los An- 
geles, xi+693 pp. 

Curran, C. H. 1932. The Norwegian zoological expe- 
dition to the Galapagos Islands 1925, conducted 
by Alf Wollebaek. IV. Diptera. (Excl. of Tipulidae 
and Culicidae). Meddelelser fra det Zoologiske 
Museum, Oslo 30: 347-366. 

1934. The Families and Genera of North 
American Diptera. The Ballou Press, New York, 
512 pp. 

Foster, G. A. 1976a. Notes on the phylogeny of the 
Nearctic Tethinidae and a review of the genus 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Neopelomyia Hendel, and the Tethina milichioides 
group (Diptera). Proceedings of the Entomologi- 
cal Society of Washington 78(3): 336-352. 

. 1976b. 74. Family Tethinidae, pp. 1—4. Jn Pa- 
pavero, N., ed., A Catalogue of the Diptera of the 
Americas South of the United States. Museu de 
Zoologia, Universidade de Sao Paulo, Sao Paulo. 

Foster, G. A. and W. N. Mathis. 1998. A revision of 
the family Tethinidae (Diptera) from the Carib- 
bean, Gulf of Mexico, and Bermuda. Proceedings 
of the Entomological Society of Washington 
100(4): 601—632. 

Hendel, E 1934. Revision der Tethiniden (Dipt. Mus- 
cid. acal.). Tijdschrift voor Entomologie 1934: 
37-54. 

Hennig, W. 1937. Systematisch-tiergeographische Bei- 
trage zur Kenntnis der Tethiniden (Dipt., Aca- 
lypt.). Entomologischen Rundschau 54(9)(1936): 
136-140. 

Loew, H. 1869. Diptera Americae septentrionalis in- 
digena. Berliner Entomologische Zeitschrift 13: 
1-52. 

Malloch, J. R. 1913. A synopsis of the genera of Agro- 
myzidae, with descriptions of new genera and spe- 
cies. Proceedings of the United States National 
Museum 46: 127-154. 

. 1934. Tethinidae, pp. 452—460. Jn Edwards, 
F, ed., Diptera of Patagonia and South Chile. Brit- 
ish Museum (Natural History), London. 

Mathis, W. N. and L. Munari. 1996. World catalog of 
the family Tethinidae (Diptera). Smithsonian Con- 
tributions to Zoology 584: 1—27. 

Melander, A. L. 1913. A synopsis of the dipterous 
groups Agromyzinae, Milichinae, Ochthiphilinae 
and Geomyzinae. Journal of the New York Ento- 
mological Society 21(4): 283-300. 

. 1952. The North American species of Teth- 
inidae (Diptera). Journal of the New York Ento- 
mological Society 59: 187-212. 

McAlpine, J. EF 1981. Morphology and terminology— 
Adults [chapter], pp. 9-63. In McAlpine, J. EF, et 
al., eds., Manual of Nearctic Diptera. Ottawa: Ag- 
riculture Canada, Research Branch, Monograph 
27, Vol. 1, 674 pp. 

Prado, A. P. do, and O. Tavares. 1966. S6bre duas es- 
pécies novas do género “‘Tethina”’ Haliday, 1838 
(Diptera, Tethinidae). Revista Brasileira de Biol- 
ogia 26(4): 429—439. 

Vockeroth, J. R. 1965. Family Tethinidae, pp. 726— 
728. In Stone, A., et al., eds., A Catalog of the 
Diptera of America North of Mexico. USDA Ag- 
ricultural Handbook 276, Washington, D.C., 
1696 pp. 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 549-560 


LIFE HISTORY OF THE PUTNAM SCALE, DIASPIDIOTUS ANCYLUS 
(PUTNAM) (HEMIPTERA: COCCOIDEA: DIASPIDIDAE) ON BLUEBERRIES 
(VACCINIUM CORYMBOSUM, ERICACEAE) IN NEW JERSEY, WITH A 
WORLD LIST OF SCALE INSECTS ON BLUEBERRIES 


SRIDHAR POLAVARAPU, JOHN A. DAVIDSON, AND DOUGLASS R. MILLER 


(SP) Blueberry and Cranberry Research Center, Rutgers University, Chatsworth, NJ 
08019, U.S.A. (e-mail: polavarapu @ aesop.rutgers.edu); (JAD) Department of Entomolo- 
gy, University of Maryland, College Park, MD 20742, U.S.A. (e-mail: 
jd12@umail.umd.edu); (DRM) Systematic Entomology Laboratory, PSI, Agricultural Re- 
search Service, USDA, Bldg. 005, BARC-W, Beltsville, MD 20705, U.S.A. (e-mail: 
dmiller @ sel.barc.usda.gov) 


Abstract.—Life history of the Putnam scale was investigated during 1997 and 1998 on 
highbush blueberries in the pine barrens of southern New Jersey. Putnam scale has two 
generations each year. Crawler emergences in the first and second generations peaked 
during late May and early to mid-August, respectively. This species overwinters as second 
instar nymphs, primarily under the bark (cork cambium) of the host. Adult females that 
occur on or under the bark of blueberries differ morphologically from those on the leaves 
and fruit. Descriptions of both forms are provided. Nine species of parasitoids were reared 
from canes containing Putnam scale infestations and peak emergence times of the para- 
sitoids coincided with the transition between the adult females and crawlers. 


Key Words: _ scale insect, life history, armored scale, Putnam scale, blueberry, pest, par- 


asitoids, lady beetles, Diaspidiotus ancylus 


Scale insects are frequently cited as pests 
of blueberries (Marucci 1966, Milholland 
and Meyer 1984, Antonelli et al. 1992), but 
there often is considerable misinformation 
about the species that are causing problems. 
Examples of erroneous statements from the 
literature include: terrapin scale, Mesole- 
canium nigrofasciatum (Pergande) secretes 
a rigid cover over its body (Milholland and 
Meyer 1984); Putnam scale overwinters as 
fully developed adults (Antonelli et al. 
1992); all scale insects on blueberries have 
a single generation each year (Milholland 
and Meyer 1984); Putnam scale secretes 
honeydew which covers leaves and fruit 
and interrupts normal plant growth (Maruc- 
ci 1966). In fact, terrapin scale does not se- 


crete a cover, but the hard, banded structure 
that is evident on the host plant is the body 
of the adult female; Putnam scale is only 
known to overwinter as second-instar males 
and females (Tinker 1957, Stimmel 1976); 
Putnam scale is reported to have two gen- 
erations each year in Delaware (Bray 
1974); armored scales do not secrete hon- 
eydew but concentrate the anal secretion 
and incorporate it into the scale cover (Fol- 
di 1989). 

The purpose of this paper is to provide 
definitive information on the life history of 
Putnam scale on blueberries in the pine bar- 
rens of southern New Jersey, to provide de- 
tailed illustrations of the leaf and stem 
forms of the species, to give information on 


550 


natural enemies reared during life-history 
studies, and to provide a list of the scale- 


insect species that occur on blueberries and 
other Vaccinium hosts. Phenological infor- 
mation on Putnam scale will enable the de- 


velopment of effective management strate- 
gies timed to coincide with the occurrence 
of susceptible stages. Data on natural ene- 
mies should assist pest management spe- 
cialists in the development of IPM pro- 
grams that do not affect natural-enemy pop- 
ulations. The list of scale-insect species on 
blueberries provides general information on 
the distribution of species and heightens 
awareness of the diverse scale fauna that 
could become pests of blueberries. 

The pest status of Putnam scale is vari- 
able. Large populations are reported to re- 
duce plant vigor (Antonelli et al. 1992). It 
also is an aesthetic pest; the fruit can be 
deformed because of depressions formed 
under aggregated females, and the scale 
covers appear as white spots on the berries. 
Feeding on leaves and green stems causes 
red areas around the feeding site. Regular 
pruning to remove older canes appears to 
keep Putnam scale populations from be- 
coming a serious problem (Weiss and Beck- 
with 1945, Marucci 1966). Application of 
dormant oil before the plant blossoms also 
is an effective control method (Marucci 
1966). 

Putnam scale life history and identifica- 
tion is complicated by the presence of dif- 
ferent morphs on different parts of the bush. 
Host-position dimorphism was first discov- 
ered on maple by Stannard (1965). He pro- 
vided evidence that the bark form was typ- 
ical of Aspidiotus (= Diaspidiotus) ancylus, 
but when bark females produced crawlers 
that settled on leaves the resulting leaf 
adults were typical of Aspidiotus howardi 
Cockerell or A. comstocki Johnson. Stan- 
nard (1965) stated that more than 90% of 
the leaf population dispersed back to the 
twigs in late summer as crawlers and im- 
plied that a significant proportion of the 
summer-generation crawlers settled on the 
leaves of the host. The bark form (Fig. 3) 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


was characterized by having the second pair 
of pygidial lobes either absent or greatly re- 
duced, and by having the interlobular plates 
with only small fimbriations. The leaf form 
(Fig. 4) has large second lobes and the 
plates have conspicuous fimbriations. These 
differences are so significant that the bark 
form was placed in the genus Diaspidiotus 
(Borchsenius 1966) and the leaf form was 
put in Abgrallaspis (Balachowsky 1953). 

A summary of the literature on the life 
history of Putnam scale is as follows. This 
species has one generation each year in 
northern areas (e.g., Iowa, parts of New Jer- 
sey, Ohio, and Pennsylvania) and two gen- 
erations in southern areas (e.g., southern II- 
linois and Delaware). Crawlers are reported 
in late spring or early summer in Iowa (Put- 
nam 1880), in May and July in Delaware 
(Bray 1974), before midsummer in Ohio 
(Houser 1918), in May or June and mid- 
summer in Illinois (Stannard 1965). Stim- 
mel (1976) states that crawlers are present 
in Pennsylvania (a one generation per year 
area) for 4—5 weeks and are active through 
late July. In Illinois, Tinker (1957) reports 
crawler peaks in the third week of June and 
again in the second week of August. The 
species overwinters as second instars on the 
bark of twigs in both single-generation 
(Stimmel 1976) and two-generation areas 
(Tinker 1957). In Illinois, Tinker (1957) re- 
ported that females lay an average of 49 
eggs at a rate of 2-3 eggs each day; eggs 
hatch in about 16 hours. Adults appear in 
May and July in Illinois (Tinker 1957) and 
in April in Pennsylvania (Stimmel 1976). 

The known parasitoids of the Putnam 
scale are as follows: Aphelinidae: Coccob- 
ius (= Phycus) varicornis (Howard); En- 
carsia (= Prospaltella) aurantii (Howard); 
E. (= Aspidiotiphagus) citrinus (Craw) 
(Gordh 1979). Tinker (1957) reared six 
‘“‘eulophid’’ species from this scale in Illi- 
nois, but the identity (and family assign- 
ment) of these species is unknown. 


METHODS 


Monitoring the life history of Putnam 
scale was undertaken using two methods. 


VOLUME 102, NUMBER 3 


Crawler emergence was monitored using 
sticky-tape traps (Scotch™ poster tape 
#109, 3M Company, St. Paul, MN) 
wrapped around infested canes as described 
by Dreistadt et al. (1994). The sticky-tape 
traps were placed on at least seven different 
plants in two different locations (near 
Browns Mills, Burlington County, New Jer- 
sey, and Rutgers Blueberry and Cranberry 
Research Center, Chatsworth, Burlington 
County, New Jersey). At the Browns Mills 
location ‘Bluecrop’ blueberries were sam- 
pled and at Chatsworth the field contained 
a mixture of mid-season varieties. Some 
plants contained two sticky-tape traps and 
each location had 10 to 12 sampling sites. 
The sticky-tape traps were replaced at about 
weekly intervals and were examined using 
a Nikon SMZ-U stereo microscope at 30— 
40 magnification. The total number of 
crawlers on each sticky-tape trap was 
counted and recorded for each sampling in- 
terval. 

The second method of life-history mon- 
itoring was undertaken by examining 
woody canes (and the leaves and fruit at 
their apex) that were pruned from plants at 
the Rutgers Blueberry and Cranberry Re- 
search Center, Chatsworth. Two sites were 
sampled; one from the northern part of the 
blueberry breeding plot near the area sam- 
pled using sticky tapes, and one from a 
southern area of the same breeding plot. 
Five canes were taken from each location 
and were examined in the laboratory using 
a Wild Photomakroskop M400 stereomicro- 
scope at 30—60X magnification. From each 
location the sex and life-history stage of the 
first 50 specimens encountered were record- 
ed; observations on parasite emergence, 
predator activity, and scale behavior were 
made during the counting process. Samples 
were taken approximately once each month. 

The following technique was used to col- 
lect the parasitoids in the samples. After 
completing the counting process, heavily 
infested pieces of blueberry cane were 
placed in clean 2 lb. 7 oz. coffee cans that 
were covered by tissue paper and held in 


551 


place by plastic lids with the centers cut 
out. Each can contained an average of 10 
twigs, five inches long, between % and 1 
inch in diameter. All parasitoids that 
emerged in the cans were collected in al- 
cohol and submitted for identification. 
Search of the literature for species of 
scale insects on Vaccinium was not exhaus- 
tive, but included the majority of the spe- 
cies on this host genus. Primary sources of 
information were: ScaleNet which is part of 
the database system within the Systematic 
Entomology Laboratory’s web site (http:// 
www.sel.barc.usda.gov), Borchsenius 
(1966), Dekle (1976), Hamon and Williams 
(1984), Kozar (1998), Howell and Kosztar- 
ab (1972), Lambdin and Kosztarab (1977). 


RESULTS 


Putnam scale life history The Putnam 
scale has two generations each year in the 
blueberry growing areas of the pine barrens 
of southern New Jersey (Fig. 1). Peak 
crawler emergences were in late May and 
early June for the first generation, and early 
to mid-August for the second (Figs. 2a, b). 
Second-instar males and females were the 
overwintering stages, and they were pre- 
dominant under the bark of old canes. 

Spring activity began in early February 
when second-instar males began to molt to 
third instars. By the end of March most sec- 
ond-instar females had transformed into 
adults (3rd instar) and males had matured 
to pupae (4th instar) and adults (5th instar) 
(Tables 1—2). In late April most of the pop- 
ulation was in the adult-female stage indi- 
cating that adult males had emerged, mated, 
and died. Crawlers (1st instars) began to ap- 
pear in mid-May and were present for the 
rest of the growing season into October. 
However, there were two distinct peaks in 
crawler abundance coinciding with the be- 
ginning of the two annual generations (Figs. 
2a, b). In early to mid-July, the number of 
crawlers taken on the sticky-tape traps de- 
creased significantly and signaled the tran- 
sition between generations. In most instanc- 
es, the sticky-tape traps at this time of year 


2nd m&f 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


3rd f 
Ca 3rd m “ios 
ea a 4th m t. oe 
oes, Sth m baie ae 


crawler peak 


Ist m&f 


crawler peak 


v 


CS Se es eee Se Sener | 


J F M A M J 


Fig. 1. 


A S O N D J 


Seasonal occurrence and duration of various instars of Diaspidiotus ancylus through the 1998 growing 


season at Rutgers Blueberry and Cranberry Research Center, Chatsworth, NJ. Points on the life-history bars are 
actual observations. Doted lines are given when direct observations of a particular instar were not made, but 
they are surmised to be present based on indirect evidence (m = male; f = female). 


were without crawlers, but a few contained 
a small number (Figs. 2a, b). Crawlers 
seemed to prefer settling under the flaky 
bark on older canes, but small numbers also 
settled on the undersides of leaves and fruit. 
As the crawlers mature, the tan cork cam- 
bium of the bark grows over the scale cov- 
er, and the only evidence that a scale insect 
is present is a rounded swelling on the bark. 
Peak emergence of crawlers at Browns 
Mills occurred a few days after peak emer- 
gence at Chatsworth, most likely because of 
slightly lower temperatures at Browns 
Mills. 

Second instars were first collected in mid 
June and were apparently present until mid- 
July (Tables 1—2). Second-instar males be- 
gan to elongate their scales in late June and 


became distinguishable from the round 
scales of the second-instar females at that 
time. We did not find sufficient numbers of 
males in the second generation to make 
good observations on their developmental 
time frame. Many empty male scale covers 
were found in samples from July 9 sug- 
gesting that adult males had already 


‘emerged. We found a few adult males on 


the sticky-tape traps from June 26. Adult 
females were first detected in mid-June, but 
scale covers were not enlarged until mid- 
July. Observations from the August 11 
sample indicate that about half of the adult 
females had eggs under their scale covers. 
The remaining females had not yet begun 
to lay eggs. Crawlers of the second gener- 
ation began to appear in late July; peak 


VOLUME 102, NUMBER 3 


500 
400 


+ SE) 


07 11 


May June 


1000 
800 


No. of Crawlers/Tape (mean 


13 26 9 25 9 


May June 


27 10 25 11 


1.3 18 4 
July 


553 


a) Chatsworth 


30°16 28 18 7 23 
July Aug. Sept. Oct. 


15 28 14 6 


Aug. Sept. Oct. 


Dates in 1998 


Fig. 2. 


Crawler abundance (mean number per sticky-tape trap + SE) of Diaspidiotus ancylus at Rutgers 


Blueberry and Cranberry Research Center, Chatsworth, NJ, (a) and at Browns Mills, NJ (b) during 1998 growing 


season. 


numbers were found on the sticky-tape 
traps in early August at the Blueberry and 
Cranberry Research Center and slightly lat- 
er at the Browns Mills location (Figs. 2a, 
b). Second instars first appeared in early 
September and by late October were the 
only stage present. In the October 30 sam- 
ple, second-instar males had begun to elon- 
gate their scale covers and were distin- 


guishable from second-instar females at that 
time. 

Host-position dimorphism.—Studies 
were undertaken to examine morphological 
differences induced by settling site loca- 
tions. Specimens collected on the stems and 
even leaf petioles showed the morphology 
typical of the ancylus form (Fig. 3) that 
have the second lobes absent or reduced to 


554 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 1. Percent of Diaspidiotus ancylus populations in each instar during the 1998 growing season at the 
north side of the blueberry breeding plot at the Rutgers Blueberry and Cranberry Research Center, Chatsworth, 


NJ. A total of 50 specimens were examined on each sampling date. Abbreviations are m = male; f = female. 
: Percent of Population in each Instar Dates in 1998 
Instars a 2 2/26 3/26 4/30 6/18 7/9 8/11 10/9 10/30 
Settled Ist 30 36 4 
Active Ist 8 22 
2nd m or f? 56 100 
2nd f 54 60 6 92 
2nd m 12 2 2, 
3rd f 56 100 6 96 42 4 
3rd m 34 8 6 2 
4th m 30 16 
Sth m 16 


small points (Fig. 3A), the plates between 
the median and second lobes have small 
fimbriations (Fig. 3B) (compared to the ho- 
wardi-comstocki form), with 18—48(31) 
macroducts on each side of the pygidium 
(Fig. 3C), 9-22(15) perivulvar pores on 
each side of the pygidium (Fig. 3D), and 
many microducts near the dorsomarginal 
area of the thorax and head (Fig. 3E). Spec- 
imens collected on fruit and leaves showed 
the morphology typical of the howardi or 
comstocki forms (Fig. 4) that have well-de- 
veloped second lobes (Fig. 4A), the plates 
between the median and second lobes have 
large fimbriations (Fig. 4B), (compared to 


Table 2. Percent of Diaspidiotus ancylus popula- 
tions in each instar during the 1998 growing season at 
the south side of the blueberry breeding plot at the 
Rutgers Blueberry and Cranberry Research Center, 
Chatsworth, NJ. A total of 50 specimens were exam- 
ined on each sampling date. Abbreviations are m = 
male; f = female. 


Percent of Population in each Instar 


Dates in 1998 

Instars 3/26 44/30 6/18 7/10 8/11 10/9 10/30 
Settled Ist 22 34522 
Active Ist 22 34 
2nd m or f? 54 78 100 
2nd f 60 
2nd m 
3rd f 100 2 OOS? 
3rd m 4 
4th m 36 
5th m 


the ancylus form), with 12—22(17) macrod- 
ucts on each side of the pygidium (Fig. 4C), 
8—10(9) perivulvar pores on each side of the 
pygidium (Fig. 4D), and few or no microd- 
ucts near the dorsomarginal area of the tho- 
rax and head (Fig. 4E). In some cases in- 
termediate forms are collected particularly 
on green stems and leaf petioles. 

There also is a striking difference in the 
appearance of the scale cover between the 
leaf-fruit form and the bark form. The ho- 
wardi-comstocki form on the fruit has a 
conspicuously white scale cover with a 
slight grayish tinge and is never under the 
outer cell layer of the host. The ancylus 
form on the bark has the scale cover dark 
gray and is usually hidden under the host 
cork cambium. 

Information on natural enemies.—Based 
on parasitoid rearing data, there were two 
occurrence peaks of adult parasitoids. The 
sample from April 30 had an emergence of 
more than 50 adult parasitoids. This time 
frame coincided with the predominance of 
adult females just before the first crawlers 
appeared. The sample from August 11 had 
an emergence of about 100 adult parasit- 
oids. The scale population at this time was 
near the end of the first generation, with 
adult females predominantly laying eggs 
and with crawlers of the second generation 
the most abundant life stage. The parasit- 
oids that were collected are as follows: 


VOLUME 102, NUMBER 3 


es 
a 


ee "eo 
ee 
ee" 


Biss3: 
C, Macroducts; D, Perivulvar pores; E, Microducts. 


Aphelinidae: Ablerus clisiocampae (Ash- 
mead); Coccobius varicornis (Howard); 
Coccophagoides sp. #1; Coccophagoides 
sp. # 2; Encarsia sp.; Encarsia aurantii 


35/5) 


Bark form (ancylus form), adult female Diaspidiotus ancylus. A, Second lobe; B, Interlobular plates; 


(Howard); Marietta carnesi (Howard). En- 
cyrtidae: Epitetracnemus intersectus (Fon- 
scolombe). Signiphoridae: Signiphora sp. 
The most abundant parasitoids were Able- 


556 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


yea 
we 


Fig. 4. Leaf and fruit form (howardi-comstocki form), adult female Diaspidiotus ancylus (Putnam). A, Sec- 
ond lobe; B, Interlobular plates; C, Macroducts; D, Perivulvar pores; E, Microducts. 


VOLUME 102, NUMBER 3 


rus clisiocampe and Marietta carnesi 
(Howard).; Coccophagoides sp. #1; Coc- 
cophagoides sp. # 2; and Coccobius vari- 
cornis were next most numerous, and the 
remainder were uncommon. 

The lady beetle (Coccinellidae) Microw- 
esia misella (LeConte) was commonly en- 
countered during the warm parts of the year, 
as was an occasional Chilocorus specimen. 
Adults were found in the samples collected 
February 2 and April 30, and larvae were 
seen in the March 26 and August 11 sam- 
ples. The predatory mite (Hemisarcopti- 
dae), Hemisarcoptes malus (Shimer) was 
noticeably abundant in the August 11 sam- 
ple. 

Survey results.—We found the following 
scale species in 1997 and 1998 in commer- 
cial blueberry fields in New Jersey: Cocci- 
dae: European fruit lecanium—Partheno- 
lecanium corni (Bouché) (4 locations); Cot- 
tony hydrangea scale—Pulvinaria hydran- 
geae Steinweden (2 locations); Cottony 
maple scale—Pulvinaria innumerabilis 
(Rathvon) (3 locations). Diaspididae: Put- 
nam scale—Diaspidiotus ancylus (6 loca- 
tions). Pseudococcidae: Blueberry mealy- 
bug—Dysmicoccus vaccini Miller and Po- 
lavarapu (7 locations). 

We collected additional species in native 
blueberries near commercial fields in NJ as 
follows: Coccidae: Cottony azalea scale— 
Pulvinaria ericicola McConnell (8 loca- 
tions). Pseudococcidae: Myrmecophile 
mealybug—FPeliococcus flaveolus (Cocker- 
ell) (7 locations). 

Other species known to occur on Vaccin- 
ium in the Northeastern US are: Cerococ- 
cidae: Cerococcus kalmiae Ferris (Eastern 
US, Kansas, Texas). Coccidae: Thorn 
scale—Eulecanium tiliae (Linnaeus) (US 
and Europe); Terrapin scale—Mesolecan- 
ium nigrofasciatum (Pergande) (eastern 
US); Cottony camellia scale—Pulvinaria 
floccifera (Westwood) (Cosmopolitan). 
Diaspididae: Cranberry scale—Abgrallas- 
pis oxycoccus (Woglum) (Eastern US); 
Oystershell scale—Lepidosaphes ulmi (Lin- 
naeus) (Cosmopolitan); San Jose scale— 


S5y/ 


Quadraspidiotus perniciosus (Comstock) 
(Cosmopolitan); Dearness scale—Rhizas- 
pidiotus dearnessi (Cockerell) (US and 
Mexico). Eriococcidae: Azalea bark scale— 
Eriococcus azaleae (Comstock) (US); Oak 
felt scale—Eriococcus quercus (Comstock) 
(US). Lecanodiaspididae: Common false pit 
scale—Lecanodiaspis prosopidis (Maskell) 
(US and Mexico). Pseudococcidae: Helio- 
coccus osborni (Sanders) (Eastern US and 
Colorado); False puto mealybug—Phena- 
coccus rubivorus Cockerell (Eastern US 
and New Mexico); Kellogg mealybug— 
Radicoccus kelloggi (Ehrhorn and Cocker- 
ell) (US). 

Other species known to occur on Vaccin- 
ium in parts of the US other than the North- 
east are: Coccidae: Barnacle scale—Cero- 
plastes cirripediformis Comstock (Cosmo- 
politan); Indian wax scale—Ceroplastes 
ceriferus (Fabricius) (Cosmopolitan); Flor- 
ida wax scale—Ceroplastes floridensis 
Comstock (Cosmopolitan); Chinese wax 
scale—Ceroplastes sinensis Del Guercio 
(Cosmopolitan); Brown soft scale—Coccus 
hesperidum Linnaeus (Cosmopolitan); Pyr- 
iform scale—Protopulvinaria pyriformis 
(Cockerell) (Tropical areas); Cottony maple 
leaf scale—Pulvinaria acericola (Walsh 
and Riley) (Eastern US); (Southern US, Pa- 
cific Islands, Caribbean Islands, Galapagos 
Islands, Israel) Urbicola soft scale—Pulvi- 
naria urbicola Cockerell (Southern US, Pa- 
cific Islands, Caribbean Islands, Galapagos 
Islands, Israel); Hemispherical scale—Sais- 
setia oleae (Olivier) (Cosmopolitan). Dias- 
pididae: Cyanophyllum scale—Abgrallas- 
pis cyanophylli (Signoret) (Cosmopolitan); 
Red bay scale—Acutaspis perseae (Com- 
stock) (New World and Europe); Oleander 
scale—Aspidiotus nerii (Bouché) (Cosmo- 
politan); Spinose scale—Aspidiotus spino- 
sus Comstock (Cosmopolitan); Camellia 
mining scale—Duplaspidiotus clavigera 
(Cockerell) (Florida, Tropical areas); Tes- 
serate scale—Duplaspidiotus tesseratus 
(Grandpré and Charmoy) (Tropical areas); 
Latania scale—Hemiberlesia lataniae (Sig- 
noret); Mimosa scale—Melanaspis mimo- 


558 

sae (Comstock) (Mexico, Arizona, Califor- 
nia, and Florida); Harper scale—Neopin- 
naspis harperi McKenzie (California, Flor- 
ida, Georgia, Hawaii, Japan, Taiwan); 
Camphor scale—Pseudaonidia duplex 


(Cockerell); White peach scale—Pseudau- 
lacaspis pentagona (Targioni-Tozzetti) 
(Cosmopolitan); False parlatoria scale— 
Pseudoparlatoria parlatoriodes (Com- 
stock) (Tropical areas); Dentate scale—Ve- 
lataspis dentata (Hoke) (Southeastern US, 
Panama). Eriococcidae: Texas eriococcin 
Eriococcus texanus (King) (Western US). 
Pseudococcidae: Bilberry mealybug—Cu- 
cullococcus vaccinii Ferris (California). Pu- 
toidae: Pacific mealybug—Puto pacificus 
McKenzie (California). 

Scale species known to occur on Vaccin- 
ium outside of the US: Coccidae: Eulecan- 
ium distinguendum (Douglas) (England); 
Eulecanium franconicum (Lindinger) (Eu- 
rope and Former Soviet Union); Partheno- 
lecanium rufulum (Cockerell) (Europe and 
Former Soviet Union); Phyllostroma myr- 
tilli (Kaltenbach) (Europe and Former So- 
viet Union); Pulvinaria peregrina (Bor- 
chsenius) (Azerbaijan and Georgia). Con- 
chaspididae: Conchaspis vaccinii Khoo 
(Malaysia). Diaspididae: Aulacaspis erica- 
cearum Takagi (Japan); Chionaspis salicis 
(Linnaeus) (Europe, Asia, N. Africa); Ni- 
veaspis vulcania Ferris (Panama); Pseudau- 
lacaspis ericacea (Ferris) (China); Quad- 
raspidiotus bavaricus (Lindinger) (Europe); 
Quadraspidiotus ostreaeformis (Curtis) 
(temperate areas); Quadraspidiotus zonatus 
(Frauenfeld) (Europe, Middle East, and Af- 
rica). Eriococcidae: Eriococcus baldonensis 
(Rasina) (Finland, Latvia, Ukraine, and 
Russia); Eriococcus costaricensis (Cocker- 
ell and Robinson) (Costa Rica); Eriococcus 
devoniensis (Green) (Europe); Eriococcus 
uvaeursi (Linnaeus) (Europe). Ortheziidae: 
Arctorthezia cataphracta (Olafsen) (Eu- 
rope); Newsteadia floccosa De Geer (Eu- 
rope). Pseudococcidae: Atrococcus bejbien- 
koi Kozar and Danzig (Hungary and For- 
mer Soviet Union); Atrococcus cracens 
Williams (Europe); Jndococcus acanthodes 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


(Wang) (China); Phenacoccus insularis 
Danzig (Russia); Phenacoccus vaccinii 
(Danzig) (Russia); Spinococcus calluneti 
(Lindinger) (Europe). Putoidae: Puto janet- 
scheki Balachowsky (France and Czechos- 
lovakia); Puto tubulifer Danzig (Mongolia 
and Russia); Puto vaccinii (Coquillett) 
(Russia). 


DISCUSSION 


It is now clear that references to the Put- 
nam scale having only a single generation 
in the commercial blueberries areas near the 
pine barrens of New Jersey are incorrect. It 
also is incorrect that the overwintering 
stage is the adult female. Quite clearly, 
there are two generations each year, and the 
overwintering stage is the second-instar 
male and female. 

Although we did not make a detailed 
comparison of the relative abundance of 
Putnam scale on the bark versus the leaves 
and fruit, it is obvious that only a very 
small proportion of the population is found 
on plant parts other than the roughened 
bark. The reasons for this are not clear, but 
there are at least two possibilities. 1) It is 
feasible that the preferred feeding site is un- 
der the bark, and only when this habitat is 
crowded will crawlers settle on sites that 
are less than optimal. 2) It also is possible 
that a large proportion of the population ac- 
tually settles on exposed parts of the host 
but is killed by pesticides used to control 
other blueberry pests. Our current thinking 
favors hypothesis 1 since we see no evi- 
dence of massive pesticide kills of crawlers 
on exposed parts of the plants, and it ap- 
pears that crawlers settle on host areas other 
than the bark only in situations where the 
bark is already encrusted with heavy pop- 
ulations of the scale. Hypothesis 1 supports 
the long-held belief that removal of old 
woody canes will prevent the Putnam scale 
from becoming a serious pest. Without the 
roughened bark of older canes to settle un- 
der, their preferred habitat is lacking, and 
the scales that settle on the smooth areas of 
the host may be more susceptible to natural 


VOLUME 102, NUMBER 3 


enemies. Our observations suggest that 
there is a much higher rate of parasitism on 
individuals that are exposed on the fruit, 
leaves, or smooth bark than on those hidden 
under rough bark. It is interesting that every 
blueberry farm that we examined for Put- 
nam scale had populations of the scale un- 
der the bark of the older canes. This sug- 
gests that these residual populations could 
build to pest levels if conditions encourag- 
ing their build up were to occur. 

Adult parasitoids seem to be most abun- 
dant when adult females and crawlers are 
most prevalent. This situation is problem- 
atic because pesticide applications (other 
than horticultural oil sprays) are most ef- 
fective against the scales during these same 
periods, i.e., when the crawlers are wander- 
ing on the host searching for settling sites 
and have yet to form a scale cover. If chem- 
ical applications are required at these times, 
it would probably be best to use horticul- 
tural oils at summer rates without combi- 
nations of pesticides, since residues from 
the latter can kill parasitoids for one or 
more weeks after application. 

A broad diversity of scale insects has 
been reported to feed on Vaccinium hosts 
including highbush blueberries. At present 
we know of 54 species of scale insects that 
feed on species of Vaccinium. In the North- 
eastern U.S. there are 21 different species. 
In other parts of the U.S. there are seven 
additional species. In areas outside of the 
U. S. there are 26 more species. Of these 
26 species, 11 have limited host ranges: Au- 
lacaspis ericacearum—Ericaceae only; Er- 
tococcus baldonensis—Ericaceae and one 
other family; E. costaricensis—Vaccinium 
only; Eulecanium distinguendum—Vaccin- 
ium only; E. franconicum—Ericaceae only; 
Indococcus acanthodes—Vaccinium only; 
Niveaspis vulcania—Vaccinium only; Phen- 
acoccus vaccinii—Ericaceae only; Phyllos- 
troma myrtilli-Ericaceae only; Puto tubu- 
lifer-Vaccinium only; Puto vaccinii—Vac- 
cinium only. If any of these species were 
accidentally introduced into the blueberry 


559 


growing areas of the U.S. there could be 
serious consequences. 


ACKNOWLEDGMENTS 


Thanks to Vera Kyryczenko, Blueberry 
and Cranberry Research Center, Rutgers 
University (BCRC) for assistance in ex- 
amining sticky-tape traps. We are grateful 
to Michael E. Schauff and Robert D. Gor- 
don, Systematic Entomology Laboratory, 
Agricultural Research Service, United 
States Department of Agriculture, Washing- 
ton, D.C. (SEL) for respectively identifying 
the Chalcidoidea and Coccinellidae report- 
ed in this manuscript. To John Bertino, Walt 
Adams, and Paul Macrie we extend special 
appreciation for allowing us to study scale 
populations on their properties. We also are 
grateful to Devon Fowler for assistance in 
collecting samples from Variety Farms, 
Hammonton, New Jersey. This research 
was partially funded by a grant from the 
New Jersey Blueberry-Cranberry Research 
Council and CSREES grant (97-34155- 
3959) awarded to SP. 

We are grateful to the following individ- 
uals for reading and commenting on the 
manuscript: James E Stimmel, Pennsylva- 
nia Department of Agriculture, Bureau of 
Plant Industry, Harrisburg; Michael E. 
Schauff and FE Christian Thompson, SEL. 


LITERATURE CITED 


Antonelli, A., E. Elsner, and C. Shanks. 1992. Arthro- 
pod management. pp. 55—75. Jn Pritts, M. P, J. E 
Hancock, and B. Strik, eds., Highbush Blueberry 
Production Guide. Northeast Regional Agricultur- 
al Engineering Service Bulletin 55, 199 pp. 

Balachowsky, A. S. 1953. Sur un Hemiberlesia CkIl. 
nouveau des montagnes du Caméroun. (Homop- 
tera; Coccoidea). [Contribution a 1’étude des Coc- 
coidea de la France d’Outre mer, 8e note.] Beitra- 
ge zur Entomologie. 3: 111—115. 

Borchsenius, N. S. 1966. A Catalogue of the Armoured 
Scale Insects (Diaspidoidea) of the World. (In 
Russian). Nauka, Moscow & Leningrad. 449 pp. 

Bray, D. FE 1974. The Fieldman’s Guide to Entomol- 
ogy. Special Bulletin, Cooperative Extension Ser- 
vice, University of Delaware, 33 pp. 

Dekle, G. W. 1976. Florida armored scale insects. Jn 
Arthropods of Florida and neighboring land areas. 
Florida Department of Agriculture and Consumer 


560 


Services, Division of Plant Industry, Vol. 3: 1— 
345. 


Dreistadt, S. H., J. K. Clark, and M. L. Flint. 1994. 


Pests of Landscape Trees and Shrubs. University 
of California, Division of Agriculture and Natural 
Resources, Publication 3359, 327 pp. 

Foldi, I. 1989. 1.1.2.4 The scale cover, pp. 43-54. In 


Rosen, D., ed., Armoured Scale Insects, Their Bi- 
ology, Natural Enemies and Control, Vol. A. El- 
sevier, Amsterdam, the Netherlands, 384 pp. 

Gordh, G. 1979. Encyrtidae, pp. 890-967. In Krom- 
bein, K. V., P. D. Hurd, D. R. Smith, and B. D. 
Burks, eds., Catalog of the Hymenoptera of Amer- 
ica North of Mexico, Vol. 1. Smithsonian Insti- 
tution Press, Washington, D.C., 1,198 pp. 

Hamon, A. B. and M. L. Williams. 1984. The soft 
scale insects of Florida (Homoptera: Coccoidea: 
Coccidae). Jn Arthropods of Florida and Neigh- 
boring Land Areas. Florida Department of Agri- 
culture & Consumer Services, Division of Plant 
Industry, Gainesville, 194 pp. 

Houser, J. S. 1918. Destructive insects affecting Ohio 
shade and forest trees. Bulletin of the Ohio Ag- 
ricultural Experiment Station 332: 165—487. 

Howell, J. O. and M. Kosztarab. 1972. Morphology 
and systematics of the adult females of the genus 
Lecanodiaspis (Homoptera: Coccoidea: Lecano- 
diaspididae). Research Division Bulletin Virginia 
Polytechnic Institute and State University, Blacks- 
burg 70: 1—248. 

Kozar, E, ed. 1998. Catalogue of Palaearctic Coccoi- 
dea. Plant Protection Institute, Hungarian Acade- 
my of Sciences, Budapest, Hungary. 526 pp. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Lambdin, P. L. and M. Kosztarab. 1977. Morphology 
and systematics of the adult females of the genus 
Cerococcus (Homoptera: Coccoidea: Cerococci- 
dae). Research Division Bulletin Virginia Poly- 
technic Institute and State University, Blacksburg 
128: 1-252. 

Marucci, P. E. 1966. Insects and their control, pp. 199— 
236. In Eck, P. and N. EF Childers, eds., Blueberry 
Culture. Rutgers University Press, New Bruns- 
wick, New Jersey, 378 pp. 

Milholland, R. D. and J. R. Meyer. 1984. Diseases and 
arthropod pests of blueberries. The North Carolina 
Agricultural Research Service, Raleigh, 33 pp. 

Putnam, J. D. 1880. Biological and other notes on Coc- 
cidae. Proceedings of the Davenport Academy of 
Natural Sciences (1879-1880) 2: 293-347. 

Stannard, L. J. 1965. Polymorphism in the Putnam’s 
scale, Aspidiotus ancylus (Homoptera: Coccoi- 
dea). Annals of the Entomological Society of 
America 58: 573-576. 

Stimmel, J. E 1976. Putnam scale, Diaspidiotus an- 
cylus (Putnam), Homoptera: Diaspididae. Regu- 
latory Horticulture 2(1): 19-20. 

Tinker, M. E. 1957. Effect of DDT on the bionomics 
of Putnam scale. Aspidiotus ancylus Putn.) (He- 
miptera, Coccidae), Ph.D. Dissertation, Entomol- 
ogy Graduate College of the University of Illinois. 

Weiss, H. B. and C. S. Beckwith. 1945. The cultivated 
blueberry industry in New Jersey, including a re- 
port on the insects of the cultivated blueberry. 
New Jersey State Department of Agriculture, Cir- 
cular 356, 23 pp. 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 561—600 


REVISION OF THE MEXICANA-GROUP OF THE CYCLOTELINE GENUS 
OZODICEROMYIA BIGOT (DIPTERA: THEREVIDAE) 


STEPHEN D. GAIMARI AND MICHAEL E. IRWIN 


University of Illinois and Illinois Natural History Survey, 1101 West Peabody Drive, 
Urbana, IL 61801, U.S.A.; (SDG) Current address: Department of Entomology, Smith- 
sonian Institution, Washington, DC 20560-0169, U.S.A. 


Abstract.—The mexicana-group of the cycloteline genus Ozodiceromyia Bigot (Diptera: 
Therevidae) is treated. The two previously described species of the group, Ozodiceromyia 
mexicana Bigot (the type species) and Ozodiceromyia argentifera (Krober), are rede- 
scribed, and two new species, Ozodiceromyia livdahli and Ozodiceromyia parargentifera, 
are described. In addition, a neotype is designated for Phycus argentifer Krober, and a 
lectotype is designated for Euphycus setosus Krober, a junior subjective synonym of Ozo- 
diceromyia mexicana. A phylogenetic hypothesis is discussed for the relationships among 
these taxa, characters and their evolution are discussed, a key to species is provided, and 


each species is diagnosed. 


Key Words: 


The genus Ozodiceromyia Bigot (Dip- 
tera: Therevidae) is distributed throughout 
North and Central America, into northern 
South America, being most speciose in the 
dry regions of the southwestern United 
States, California, and northern Mexico. 
The genus was treated by Gaimari (1998) 
from the standpoint of hypothesizing rela- 
tionships among described species. As a re- 
sult of cladistic phylogenetic analysis, a 
well characterized “‘mexicana-group,”’ in- 
cluding Ozodiceromyia mexicana Bigot and 
Ozodiceromyia argentifera (Kréber), was 
demonstrated. 

The concept of Ozodiceromyia was intro- 
duced by Bigot (1890), for the single spe- 
cies Ozodiceromyia mexicana, and the ge- 
nus remained monotypic until Phycus ar- 
gentifer Krober was included by Cole 
(1965). Irwin and Lyneborg (1981la) were 
the first to recognize the diversity of Ozo- 
diceromyia, transferring 34 species into the 
genus, mostly from Psilocephala Zetter- 


phylogeny, revision, morphology, cladistics 


stedt and Thereva Latreille. The list was 
modified by Gaimari and Irwin (2000) to 
reflect current opinions on synonymy and 
to add several new combinations, leaving 
the genus with 27 valid species. The current 
revision treats the nominal mexicana-spe- 
cies group, seeking to clarify the identities 
of the two described species within the 
group, which have often been confused, and 
to add two new species, both of which can 
be confused with the described species. 


METHODS 


Specimens were examined using a Wild/ 
Leica MZ8 binocular dissecting microscope 
with a range of magnification between 6.3 
and 50. For phylogenetic analyses, a min- 
imum of four male and four female speci- 
mens were examined for each species, with 
the exception of Ozodiceromyia parargen- 
tifera, which is known from only the ho- 
lotype male and three female paratypes. 

For each species, at least one male ab- 


562 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
Table 1. Data matrix used in analysis of the Ozodiceromyia mexicana-group. 
Ozodiceromyia signatipennis 00000 00000 00000 00000 00000 00000 00000 00000 O 
Ozodiceromyia costali 01010 00001 10000 00000 10000 00010 11001 00000 O 
Ozodiceromyia argentifera 11111 11101 01101 10001 11100 10101 11110 00011 1 
Ozodiceromyia livdahli 11111 11101 01101 10001 11100 00101 11110 00011 1 
Ozodiceromyia mexicana 11011 11111 10011 11001 10111 01010 11111 110100 
Ozodiceromyia parargentifera 11111 11101 00000 10111 11000 10101 11110 OO111 1 


i 


domen and one female abdomen were mac- 
erated and dissected per the procedure out- 
lined in Gaimari and Irwin (2000). Illustra- 
tions (carbon dust on clayboard) were made 
with a camera lucida attached to a Leica 
MZ12 binocular dissecting microscope with 
a range of magnification between 8 and 
100. Morphological terminology follows 
that of Gaimari and Irwin (2000). 

Each specimen has been assigned a 
unique 6-digit number with the prefix 
‘‘MEI.”’ This number is printed on a yellow 
label (THEREVIDAE/M. E. Irwin/Speci- 
men #), which is attached to each pinned 
specimen. This unique number for each 
specimen facilitates entry and manipulation 
of data in a specimen-level database of 
world Therevidae. These numbers are re- 
ferred to throughout the text, illustrations, 
and in the ‘Materials examined” list for 
each species. 

Cladistic analysis.—Phylogenetic analy- 
ses follow the cladistic philosophy of par- 
simony promoted by Hennig (1966) and lat- 
er by other workers (e.g., Wiley 1981; Far- 
ris 1983). The same basic principles, phi- 
losophies, and specific methods are 
followed as outlined in Gaimari and Irwin 
(2000). 

Character polarities were determined us- 
ing outgroups (Watrous and Wheeler 1981; 
Farris 1982; Maddison et al. 1984; Nixon 
and Carpenter 1993) chosen within the con- 
text of the phylogeny of the described spe- 
cies of the genus proposed by Gaimari 
(1998). Cladistic analyses were performed 
using the exhaustive search option in PAUP 
(Swofford 1993, version 3.1.1) using the 
data matrix presented in Table 1. The clad- 
ogram figure displaying character state 


changes was prepared using WINCLADA 
(Nixon 1999, currently in version 0.9.9 
beta) under ACCTRAN character optimi- 
zation (Swofford and Maddison 1987), fol- 
lowing the same reasoning as Gaimari and 
Irwin (2000). 

Terminal taxa.—Very few specific hy- 
potheses exist for relationships within Ozo- 
diceromyia, and only Gaimari (1998) con- 
sidered the genus in the broader context of 
phylogeny. Nonetheless, many undescribed 
species can be assigned to species-groups 
based on putative synapomorphies. Gaimari 
(1998) suggested a close relationship be- 
tween Ozodiceromyia argentifera and Ozo- 
diceromyia mexicana based upon the fol- 
lowing synapomorphic characteristics: an- 
tenna longer than head; setae covering more 
than the basal half of first flagellomere; first 
flagellomere fully surrounded by setae 
equally on all sides; antennal style subapi- 
cal and not extending beyond tip of first 
flagellomere; halter yellow (although this 
state appears in other members of the ge- 
nus); distal part of ventral lobe of male gen- 
italia with a pilose distal secondary lobe. 
Based upon these synapomorphic states, 
two new species have been recognized as 
members of this group and are described 
herein, along with redescriptions of the pre- 
viously described species, and a phyloge- 
netic hypothesis is proposed for the rela- 
tionships among these taxa. 

The two outgroup taxa, Ozodiceromyia 
costalis (Loew, 1869) (MEI 033251, 
037735, 037741, 041847, 044821, 044990, 
050648, 075851) and Ozodiceromyia sig- 
natipennis (Cole 1923) (MEI 035882, 
041854, 041855, 043837, 044750, 056972, 
056973, 071403), were chosen on the basis 


VOLUME 102, NUMBER 3 


of relationships within this genus hypothe- 
sized by Gaimari (1998). Ozodiceromyia 
costalis is hypothesized to be closely relat- 
ed to the mexicana-group based upon the 
following synapomorphic characteristics: 
medial surface of scape setose; medial sur- 
face of pedicel setose; setae of male frons 
in patches or absent, not scattered; face di- 
rectly below antenna shiny black, with little 
or no pruinescence (however, Ozodicero- 
myia argentifera displays the state of hav- 
ing silver pruinescence, and so this state is 
only synapomorphic under accelarated 
transformation character optimization); 
presence of projection on gonocoxal apo- 
deme of male genitalia for articulation with 
aedeagus. Ozodiceromyia signatipennis is 
hypothesized to be a more distant part of 
this larger clade, based upon the following 
synapomorphic characteristics: median oc- 
cipital sclerite of male entirely shiny black, 
lacking pruinescence; ventral gonocoxal 
process of male genitalia present as a small 
fold of the edge. 


DESCRIPTIONS AND REDESCRIPTIONS OF 
SPECIES 


The recognized species of the Ozodicer- 
omyia mexicana-group are described or re- 
described herein, and a dichotomous key is 
included. An exhaustive list of references 
for the two previously described taxa is pro- 
vided by Gaimari (1998), but only taxo- 
nomically relevant references are listed 
herein. Within species descriptions, values 
in [ ] represent the values taken from par- 
atypes or additional specimens of the same 
sex. Abbreviations for repositories follow 
Arnett et al. (1993), and are listed in Table 2. 


Ozodiceromyia Bigot 


Ozodiceromyia Bigot 1890: 323. Type spe- 
cies: Ozodiceromyia mexicana Bigot, by 
original designation. Sabrosky 1978: 143 
(formally fixed spelling as “‘first revis- 
er’); Gaimari and Irwin 2000 (in phylog- 
eny, key ref., lit. list, list spp., biogeogr.). 

Ozodiceromya (incorrect original spelling): 
Bigot 1890: 321 (orig. descr.); Irwin and 


563 


Lyneborg 1981a: 203 (key ref.), 254 (re- 
descr) 198th: 522 r(key-ret:): 
Ozodiceronyma (misspelling): Godman 
1901: 378 (listing). 
Phycus Walker 1850, in part: Becker 1912: 
294 (prop. Ozodiceromyia as junior syn.). 
Psilocephala ‘‘haemorrhoidalis-group”’: 
Cole 1923: 34—37 (key ref.), 37 (descr.). 


Ozodiceromyia mexicana-group 


The small, easily recognized mexicana- 
group is characterized by an elongated an- 
tennal scape and first flagellomere, as well 
as the slightly elongated, barrel-shaped ped- 
icel. In total, the antenna is longer than the 
head, and is densely setose, including setae 
on the median surface. This surface is bare 
in nearly all other members of the genus. 
Setae cover all but the distal third of the 
first flagellomere. The stylus is inserted 
subapically in a ventral pit slightly behind 
the apex of the first flagellomere. In the 
wings, the veins are orange, at least in the 
basal two-thirds; the distal third is often 
darkened and smoky, with correspondingly 
darker veins. In the male genitalia, the gon- 
ostylus is distinctly expanded ventrally into 
a large lobe, which is densely covered with 
fine, pale or orange setae. The sclerotized 
portions of the ventral lobes are completely 
separated, and each has a small, secondary 
lobe distally, covered with fine, short pile. 
When present, the ventral gonocoxal pro- 
cess is flattened and elongated (the ventral 
gonocoxal process is absent in Ozodicero- 
myia mexicana). The outer gonocoxal pro- 
cess is flattened and elongated. In the fe- 
male terminalia, the furca lacks an anterior, 
sclerotized transverse bar. The common 
spermathecal duct originates on the furcal 
membrane within an indented cavity on the 
posterior part of the furcal bulla. 

Synapomorphies.—Antenna longer than 
head (character 1), setae of first flagellom- 
ere covering more than basal half (character 
5), setae fully surrounding first flagellomere 
(character 6), antennal style distinctly sub- 
apical (character 7), first stylar segment re- 
duced to flattened ring (character 8), wing 


564 


Table 2. 


AMNH 
ANIC 
ANSP 
ASUT 
BMNH 
BPBM 
BYUC 
CASC 
CDFA 


CICESE 


CMNH 
CNCI 
CSUC 


CUIC 
DEIC 
DENH 
EBCC 


EMEC 
EMUS 
ESUW 


FMNH 
FSCA 
IEXA 
INBC 
INHS 
IZAS 
IZAV 


KSUC 
KUIC 
LACM 
MCZC 
MEIC 
MEUC 
MHNG 
MNHN 
MRSN 
MTEC 
MUSM 
MZSP 
NHMW 
NHRM 
NMSA 
NMSU 
NVDA 
NYSM 
OSEC 
OSUC 
PMNH 
QCAZ 
SDGC 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


List of codons used for repositories of specimens. 


American Museum of Natural History, New York, New York. 

Australian National Insect Collection, CSIRO, Canberra, ACT, Australia. 

Academy of Natural Sciences, Philadelphia, Pennsylvania. 

Frank M. Hasbrouk Insect Collection, Arizona State University, Tempe, Arizona. 

The Natural History Museum, London, United Kingdom. 

Bernice P. Bishop Museum, Honolulu, Hawaii. 

Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah. 

California Academy of Sciences, San Francisco, California. 

California State Collection of Arthropods, California Department of Food and Agricul- 
ture, Sacramento, California. 

Centro de Investicacién Cientifica y de Educacion Superior de Ensenada, Baja Califor- 
nia Norte, Mexico. 

Carnegie Museum of Natural History, Pittsburgh, Pennsylvania. 

Canadian National Collection of Insects, Ottawa, Ontario, Canada. 

C. P. Gillette Arthropod Biodiversity Museum, Colorado State University, Fort Col- 
lins, Colorado. 

Cornell University Insect Collection, Ithaca, New York. 

Deutsches Entomologisches Institut, Eberswalde, Germany. 

Entomological Museum, University of New Hampshire, Durham, New Hampshire. 

Estacion de Biologia Chamela, Universidade Nacional: Autonoma de México, Jalisco, 
Mexico. 

Essig Museum of Entomology, University of California, Berkeley, California. 

Entomological Museum, Utah State University, Logan, Utah. 

Rocky Mountain Systematic Entomology Laboratory, University of Wyoming, Lara- 
mie, Wyoming. 

Field Museum of Natural History, Chicago, Illinois. 

Florida State Collection of Arthropods, Gainesville, Florida. 

Instituto de Ecologia, Xalapa, Vera Cruz, Mexico. 

Instituto Nacional de Biodiversidad, Santo Domingo, Heredia, Costa Rica. 

Illinois Natural History Survey, Champaign, Illinois. 

Institute of Zoology, Academica Sinica, Beijing, China. 

Instituto de Zoologia Agricola, Universidad Central de Venezuela, Maracay, Venezue- 
la. 

Kansas State University Insect Collection, Manhattan, Kansas. 

Kagoshima University Insect Collection, Kagoshima, Japan. 

Los Angeles County Museum of Natural History, Los Angeles, California. 

Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts. 

Michael E. Irwin private collection, for eventual deposit in CASC. 

Museo Entomologico, Universidad de Chile, Santiago, Chile. 

Muséum d’ Histoire Naturelle, Genéve, Switzerland. 

Muséum National d’Histoire Naturelle, Paris, France. 

Spinola Collection, Museu Regionale di Scienze Naturali, Torino (= Turin), Italy. 

Montana State University Entomological Collection, Bozeman, Montana. 

Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru. 

Museu de Zoologia, Universidade de Sao Paulo, Sao Paulo, Brazil. 

Naturhistorisches Museum Wien, Wien (= Vienna), Austria. 

Naturhistoriska Riksmuseet, Stockholm, Sweden. 

Natal Museum, Peitermaritzburg, Natal, South Africa. 

Insect Collection, New Mexico State University, Las Cruces, New Mexico. 

Nevada State Department of Agriculture, Reno, Nevada. 

New York State Museum, Albany, New York. 

K. C. Emerson Museum, Oklahoma State University, Stillwater, Oklahoma. 

Museum of Biological Diversity, Ohio State University, Columbus, Ohio. 

Peabody Museum of Natural History, Yale University, New Haven, Connecticut. 

Quito Catholic Zoology Museum, Universidad Catolica del Ecuador, Quito, Ecuador. 

Stephen D. Gaimari private collection. 


VOLUME 102, NUMBER 3 


Table 2. Continued. 


565 


SDMC 
SEMC 
SWRS 
Arizona. 
TAMU 
TAUI 
UAIC 
UCDC 
UCMC 
UCRC 
erside, California. 
UGCA 
UMRM 
UNAM 
Distrito Federal, México. 
UNSM 
USNM 
Columbia. 
WFBM 
WSUC 
Washington. 
ZMAS 
ZMHB 
ZMUC 


San Diego Natural History Museum, San Diego, California. 
Snow Entomological Collection, University of Kansas, Lawrence, Kansas. 
Southwestern Research Station of the American Museum of Natural History, Portal, 


Texas A&M University Insect Collection, College Station, Texas. 

Zoological Museum, Tel Aviv University, Tel Aviv, Israel. 

University of Arizona Insect Collection, Tucson, Arizona. 

Bohart Museum of Entomology, University of California, Davis, California. 
University of Colorado Museum, Boulder, Colorado. 

UCR Entomological Research and Teaching Collection, University of California, Riv- 


Museum of Natural History, University of Georgia, Athens, Georgia. 
W.R. Enns Entomology Museum, University of Missouri, Columbia, Missouri. 
Coleccion Nacional de Insectos, Universidad Nacional Aut6noma de México, México, 


University of Nebraska State Museum, Lincoln, Nebraska. 
National Museum of Natural History, Smithsonian Institution, Washington, District of 


W. F. Barr Entomological Collection, University of Idaho, Moscow, Idaho. 
Maurice T. James Entomological Collection, Washington State University, Pullman, 


Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia. 
Museum fiir Naturkunke, Humboldt Universtitat, Berlin, Germany. 
Zoological Museum, University of Copenhagen, Copenhagen, Denmark. 


cloudy yellow or orange and darkened dis- 
tally (character 16), distiphallus longer than 
dorsal apodemes (character 20), distiphallus 
recurved before apex, becoming parallel 
with dorsal apodemes (character 23; with 
subsequent change to perpendicular in Ozo- 
diceromyia parargentifera), secondary dis- 
tal lobe present at tip of ventral lobe (char- 
acter 33), ventral part of gonostylus ex- 
panded into large lobe (character 34), an- 
terior margin of furcal bulla indented as a 
cavity to house the common spermathecal 
duct (character 39). Synapomorphies for 
Ozodiceromyia parargentifera + Ozodicer- 
omyia argentifera + Ozodiceromyia livdah- 
li: setae of scape shorter than or subequal 
to setae of first flagellomere (character 3), 
basal portion of distiphallus swollen (char- 
acter 22), gonocoxal setae all black (char- 
acter 26; with subsequent change to pale 
setae in Ozodiceromyia livdahli), ventral 
gonocoxal process present, elongated, and 
flanged (character 28), setae of outer gon- 
ocoxal process present (character 30), sub- 
apical spur of gonostylus absent (character 


35), basal part of common spermathecal 
duct distinctly widened, tapering quickly 
(character 40), spermathecal ducts originate 
from common spermathecal duct (character 
41). Synapomorphies for Ozodiceromyia 
argentifera + Ozodiceromyia livdahli: 3 
notum with thick, decumbent pile (charac- 
ter 12), fine, erect setae of d notum entirely 
pale or gold (character 13). 


Ozodiceromyia argentifera (Kroéber) 
(Figs. 7-8, 13, 14-15, 17-19, 26—27, 32- 
33,739) 


Phycus argentifer Krober 1929: 418. Type 
locality: Oaxaca, Mexico. Type: NTd 
(designated herein) in ZHMB. Kroéber 
1929: 418 (comment on affinity with 
Ozodiceromyia, illust.: antenna). 

Ozodiceromyia argentifera: Cole 1965: 349 
(comb. change, cat. cit.); Gaimari 1998: 
117 (it. list), 157-159 Gn phylogeny), 
188—267 (illust.: d and @ heads, lat. tho- 
rax, wing base, ¢ and & pregenit. segs. 
and genit.); Gaimari and Irwin 2000 Gn 
phylogeny; illust.: ¢d and 2 heads, lat. 


566 


thorax, wing base, d and @ pregenit. 
segs. and genit.) 

Ozodiceromya (sic) argentifera: Irwin and 
Lyneborg 1981a: 257 (listing). 


Male.—Body length: 8.5 mm. 

Head (Fig. 7): 1.2 mm long, 2.2 mm 
wide, 1.7 mm high. Distance between eyes 
at antennal level 0.50 mm;-at genal level 
0.78 mm. Antenna brown, except basal half 
of scape orange. Scape 1.11 mm long, 0.15 
mm wide; with short, fine, black setae, 
evenly distributed over entire scape (includ- 
ing median surface), and few larger setae in 
basal half. Pedicel 0.24 mm long, barrel- 
shaped, setose (including median surface). 
First flagellomere 1.02 mm long, 0.17 mm 
wide; with short setae covering basal two- 
thirds. Stylus 0.09 mm long, inserted sub- 
apically in ventral pit. Antennal base to 
nearest edge of eye 0.18 mm. Frons bulging 
0.09 mm beyond eye in lateral view. Face 
below antenna with silver pruinescence. 
Parafacial lacking pile, with silver prui- 
nescence extending to antennal base and 
dorsally along eye margin halfway up frons. 
Frons otherwise shiny black; with few, 
short, fine, black setae (0.15 mm long) dor- 
solateral to antennal base; remainder of 
frons bare. Genal pile short, darkened. Pal- 
pal pile white basally, dark brown distally. 
Postgenal and occipital pile white; occiput 
with silver pruinescence only along edge of 
eye, and with several black setae. Median 
occipital sclerite flattened; glabrous, shiny 
black; upper edge not rounded. Postocular 
setae black, arranged in single, transverse 
row. Ocellar tubercle with silver pruinesc- 
ence, and fine, forward-directed, black se- 
tae. 

Thorax (Fig. 14): Scutum and scutellum 
with appressed and semi-appressed gold 
pile. Scutum 2.6 mm long, 1.7 mm wide; 
ground color black; with silver-blue prui- 
nescence dorsally, lacking pruinescence lat- 
erally; median vitta diffuse bronze; dc vittae 
absent; 1 pair [or lacking] dc setae. Scutel- 
lum with silver pruinescence, reduced prui- 
nescence anteriorly. Halter yellow, except 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


base of stalk brown. Katatergite with dense, 
long, white pile. Anepimeron, katepimeron, 
and meron lacking pile, and with reduced 
silver pruinescence, appearing as vertical 
brown stripe from wing base to between 
second and third coxae. Anepisternum with 
silver pruinescence, and with white pile. 
Katepisternum with silver pruinescence, 
with white pile only on vertical crest along 
middle of pleurite. Prepimeron with silver 
pruinescence, lacking pile. 

Legs: Coxae with silver pruinescence, 
posterior surfaces less so. Posterior surface 
of hindcoxa lacking pile (Fig. 14). Femora 
dark brown; with appressed, scale-like 
brown and white pile dorsally. Fore- and 
midfemora also with erect and recumbent 
brown and white pile. Tibiae with short, 
erect, black setae; proximal, dorsal surface 
with sparser setal covering. Foretibia or- 
ange on basal half, becoming dark brown 
distally; clavate distally. Midtibia orange, 
becoming darkened distally. Hindtibia or- 
ange. Foretarsus dark brown. Mid- and 
hindtarsi dark brown except for basal two 
tarsomeres mostly orange. 

Wing: 6.5 mm long. Basal costal lobe 
with setae arranged in line along outer 
edge, extending into outer row of costal se- 
tae; second line of setae along base of basal 
costal lobe, extending into inner row of cos- 
tal setae (Fig. 15). Entire membrane slightly 
darkened; veins and membrane orange ba- 
sally, brown distally. 

Abdomen: Tergites with erect and re- 
cumbent, white pile, except tergite 1 with 
recumbent brown pile mediodorsally; dor- 
sally, with silver pruinescence; laterally, 
lacking pruinescence (showing dark brown 
ground color), except tergite 1 and posterior 
edges of basal tergites with silver pruinesc- 
ence. Sternites with silver pruinescence 
only along anterior edge of sternite 2. 

Terminalia: (MEI 037789). Sternite 8 
(Fig. 17) with black setae restricted to pos- 
terior edge, which is emarginate medially. 
Tergite 8 (Fig. 18) dumbell-shaped, with 
black setae restricted to posterior edge. 
Epandrium (Fig. 23) 0.24 mm long, 0.72 


VOLUME 102, NUMBER 3 


mm wide at widest point; orange; emargin- 
ate anteriorly; dorsal surface with black se- 
tae on posterior half; lateral edges parallel; 
posterolateral corners extended posteriorly. 
Subepandrial plate (Fig. 23) attached to 
posterolateral and posterior edges; sclero- 
tized portion V-shaped posteriorly. Sclero- 
tized portion of cerci 0.24 mm long; ex- 
tending posteriorly beyond posterolateral 
corners of epandrium; subequal in length to 
ventral epandrial sclerite. Gonocoxites 
(Figs. 26-27) 0.83 mm wide; orange; with 
black setae, up to 0.45 mm long; fusion 
0.39 mm long at midline, lacking suture. 
Inner gonocoxal process flanged; knob with 
several black setae. Outer gonocoxal pro- 
cess flange-like, 0.38 mm long; with small 
patch of short, fine setae dorsodistally. Gon- 
ocoxal apodeme entirely within anterior 
edge of gonocoxite; lacking sclerotized 
bridge to parameral sheath of phallus. Ven- 
tral gonocoxal process 0.26 mm long, ta- 
pering distally; bare. Ventral lobes distinct, 
fused basally by thin, transparent mem- 
brane. Gonostylus (Fig. 19) expanded ven- 
trally into large lobe; with dorsobasal lobe; 
with subapical, lateral spur. Dorsal apode- 
mes (Fig. 32) of aedeagus parallel; subequal 
in length to ventral apodeme; parameral 
sheath smooth dorsally. Ventral apodeme 
uniformly wide; lacking ventral keel. Ejac- 
ulatory apodeme 0.38 mm long; stick-like, 
slightly expanded distally. Lateral ejacula- 
tory process a complete ring dorsally, but 
notched; set into aedeagus; lightly sclero- 
tized. Distiphallus swollen basally, long, re- 
curved; distally parallel with ventral apo- 
deme (Fig. 33). 

Female.—Similar to d except as follows: 
body length 10.0 mm. 

Head (Fig. 8): 1.4 mm long, 2.5 mm 
wide, 1.6 mm high. Distance between eyes 
at level of anterior ocellus 0.53 mm; at an- 
tennal level 1.01 mm; at genal level 1.13 
mm. Scape 1.35 mm long, 0.18 mm wide. 
First flagellomere 1.19 mm long, 0.18 mm 
wide. Antennal base to nearest edge of eye 
0.35 mm. Frons bulging 0.17 mm beyond 
eye in lateral view. Parafacial pruinescence 


567 


ends at antennal level. Frons shiny black; 
lower frons bulging to antennal base; upper 
frons bulging, distinct from lower frons, 
with short, black setae (up to 0.18 mm 
long) in patch between edge of eye and 
midline. Median occipital sclerite and upper 
edge rounded; transverse row of black setae 
across median occipital sclerite in addition 
to row of postocular setae. 

Thorax: Scutum and scutellum with 
short, recumbent, black and gold pile. Scu- 
tum 2.8 mm long, 2.1 mm wide. Pile of 
anepisternum and katepisternum short, 
white. 

Wing: 7.4 mm long. 

Abdomen: Tergites and sternites lacking 
pruinescence (showing dark brown ground 
color), except posterolateral edges of ter- 
gites 2 and 3 with silver pruinescence. Ter- 
gites covered with short, fine, appressed 
gold pile. 

Terminalia (Fig. 39): Furca 0.53 mm 
long, 0.33 mm wide; anterior edge not 
sclerotized; anterolateral prongs dorsoven- 
trally flattened, separation subequal to 
greatest width of furca. Furcal bulla not 
sclerotized; posteriorly with indented cavi- 
ty. Gonopore basal to furcal bulla, originat- 
ing within posterior cavity. Common sper- 
mathecal duct basally 0.11 mm wide, ta- 
pering to 0.05 mm within 0.33 mm distance 
from gonopore; 0.68 mm long. Spermathe- 
cal ducts originate from clean trifurcation 
with central sac duct; central sac duct wider 
in basal diameter than spermathecal ducts. 
Central sac duct 0.45 mm long. Central sac 
0.69 mm long, 0.39 mm wide. Spermatheca 
0.18 mm in diameter; rounded, but basal 
edge slightly flattened. 

Type material—NEOTYPE (here desig- 
nated): pinned 3d (MEI 027042) with the 
following labels: 10 mi. NE Haujuapan de 
Leon, Oax.(aca), Mex.(ico), VI-27-1965, 
Burke, Meyer, Schaffner/Neotypus 6, Phy- 
cus argentifer Kréber, designated in 1998 
by SD Gaimari et ME Irwin (red label)/ 
Ozodiceromyia argentifera (Kréber), det. S. 
D. Gaimari, 1997. This pinned specimen is 


568 


in excellent condition, and is deposited in 
ZMHB (with permission of TAMU). 

The designation of a neotype in this cir- 
cumstance is warranted under article 75.3 
of the International Code of Zoological No- 
menclature, Fourth Edition, for the follow- 
ing reasons: 75.3.1) designation of this neo- 
type is done for the express purpose of fix- 
ing the identity of a species where three 
similar species occur, two of which are new 
to science; 75.3.2) the characters that dif- 
ferentiate this taxon from other taxa are 
contained in both the species diagnosis and 
key in the present work; 75.3.4) the holo- 
type was confirmed lost in the postal system 
(the package was shipped from Berlin by 
post, and a broken (likely crushed open or 
deliberately slit open along the box’s edge) 
box half full with packing material arrived 
in Illinois; staff in Berlin and in Illinois 
made inquiries with the postal services, 
confirming that the specimen was lost); 
75.3.5) the original type locality (Veracruz, 
Mexico) allows only for certain possible 
identities, all of which are excluded except 
the current species due to morphological 
characteristics in the original description 
(e.g., area under antennae is silver-white ex- 
cludes Ozodiceromyia mexicana, and the 
yellow halter excludes Ozodiceromyia par- 
argentifera); the original description fully 
matches the holotype description; 75.3.6) 
Oaxaca, Mexico is as close to Veracruz as 
any of the known specimens. 

Materials examined.—MEXICO: Chi- 
huahua, Cuiteco, T. A. Sears, P C. Gardner, 
C. S. Glaser, 29-VII-1969 (1 36, MEI 
052225, UCDC); Durango, 19.3 km W of 
Dolores Hidalgo, E. M. Fisher, J. L. Fisher, 
7-I[X-1970 (1 36, MEI 037789, SDGC); 8.1 
km W of Durango, 1981.2 m, J. E Mc- 
Alpine, 23-VII-1964 (1 2, MEI 052215, 
CNCI); 11.3 km W of Durango, 2,133.6 m, 
W. C. McGuffin, 26-VII-1964 (1 2, MEI 
052230, SDGC), W. R. M. Mason, 11-VIII- 
1964 (1 2, MEI 052210, CNCI), 22-VII- 
1964 (1 2, MEI 052214, CNCI; 1 2, MEI 
052217, ZMHB), 2,286 m, W. R. M. Ma- 
son, 14-VII-1964 (1 6, MEI 052213, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


BMNH); 17.7 km W of Durango, 2,133.6 
m, J. EK McAlpine, 29-VII-1964 (1 2, MEI 
052218, CNCI), L. A. Kelton, 2-VII-1964 
(1 36, MEI 038785, CNCI); 28.98 km W of 
Durango, 2,194.56 m, J. A. Chemsak, 31- 
VII-1964 (1 3, MEI 045406, CASC); 32.2 
km W of Durango, 2,133.6 m, Rio Chico, 
J. E McAlpine, 10-VIII-1964 (1 2, MEI 
052208, CNCI), 22-VII-1964 (1 36, MEI 
052212;' 1. 2y.MEI 052216, ZMUCG ile 
MEI 052219, CNCI; 1 2, MEI 052211, 
BMNH; 1 &, MEI 052209, MEIC), 
2,194.56 m, L. A. Kelton, 23-VII-1964 (1 
3, MEI 052207, MEIC); Jalisco, 13 km. N 
Autlan [de Navarro], Carn. Mina San Fran- 
cisco, EF A. Noguera, A. Rodriguez, 29-VI- 
1995 (1 2, MEI 103438, UNAM); 13 km 
NE San Gabriel, A. Rodriguez, E A. No- 
guera, 8-X-1994 (1 6, MEI 051278, 1 °, 
MEI 051277, UNAM; 1 2, MEI 051276, 
MEIC); 16 km N of Autlan [de Navarro], 
Carroll, Friedlander, J. C. Schaffner, 7-VII- 
1984 (1 2, MEI 027077, TAMU); 40.3 km 
SE of Atotonilco, J. W. MacSwain, 23-VII- 
1952 (1 6, MEI 038789, 3 & ¢ in copula, 
MEI 052171, 052172, EMEC; 1 3, MEI 
038790, SDGC; 1 2, MEI 038788, 
AMNBH); [Estacion de] Biol.[ogia] Chame- 
la, E. Ramirez, 8-VII-1988 (1 9°, MEI 
051279, EBCC); Estacion de Biologia Cha- 
mela, E. Ramirez, 17-VII-1981 (1 6, MEI 
051280, EBCC); Guadalajara, McConnell, 
1909 (1 2, MEI 078359, CMNH); Michoa- 
can, 29 km NW of Quiroga, 2,072.64 m, 
R. H. Painter, E. M. Painter, 22-VIII-1962 
(1 2, MEI 052183, KSUC); Morelos, 7.1 
km E of Cuernavaca, Clark, Murray, Ashe, 
J. C. Schaffner, 6-VII-1974 to 7-VH-1974, 
at light, (1 2, MEI 026699, TAMU); 38 km 
W Iguala Gro. [probably referring to Iguala 
de la Indepencia, near the Morelos border 
in Guerrero], J. Chemsak, A. Michelbacher, 
M. Michelbacher, 23-VII-1983 (1 2, MEI 
081501, USNM); Nayarit, Ahuacatlan, P. 
D. Hurd, 18-VII-1951 to 22-VII-1951, on 
fllower]s. of Donnellsmithia hintonii (1 6, 
MEI 038786, AMNH); Oaxaca, 16.1 km 
NE of Huajuapan de Leon, Burke, Meyer, 
J. C. Schaffner, 27-VI-1965 (1 2, MEI 


VOLUME 102, NUMBER 3 


569 


Fig. 1. 


027043, SDGC); Puebla, 6.4 km NW Te- 
panco de Lopez, 2-VII-1952, Univ. Kans. 
Mex. Expedition (1 2, MEI 052179, 
SEMC); 8 km S of Tecomachalco, 2103.12 
m, M. E. Irwin, 10-VIII-1967, flight trap, 
narrow canyon (1 36, MEI 052195, UCRC); 
Zacatecas, 15 km E of Sombrerete, P. D. 
Hurd, 28-VII-1951 to 31-VII-1951 (1 2, 
MEI 038783, EMEC). 

Diagnosis.—The scape in this species is 
setose, but most of the setae are short, with 
few longer ones. The lower frons bulges 
only slightly, with silver pruinescence lat- 
erally, extending down along parafacial. 
The face below the antenna also has silver 
pruinescence. In males, the notum is dense- 
ly covered with erect and appressed gold 
pile. In females, the gold pile on the notum 
and abdominal tergites is short, fine, and 
appressed. In the male genitalia, the setae 
of the epandrium and gonocoxites are 
black. The subepandrial plate is attached to 
the epandrium at the posterolateral corners, 
and the sclerotized portion of the subepan- 
drial plate is V-shaped posteriorly. The gon- 
ocoxites lack a suture along the midline 
separating the two lateral halves. The outer 
gonocoxal process is widened distally, with 


Known distribution for Ozodiceromyia argentifera. 


a small patch of setae on the dorsolateral 
surface. The ventral gonocoxal process of 
the gonocoxites is present, elongated and 
flattened; there are no clumped setae at its 
base. The base of the ventral lobes is U- 
shaped and broad between the two halves. 
The parameral sheath of the aedeagus is 
smooth dorsally. The ventral apodeme lacks 
a ventral keel. The distiphallus is swollen 
basally, and is recurved and parallel with 
the ventral apodeme at the tip. In the female 
terminalia, the furca is not notched poste- 
riorly, and lacks a posterolateral peg. The 
anterolateral furcal prongs are separated by 
a distance subequal to the greatest width of 
the furca. The common spermathecal duct 
is broad basally, at half the width of the 
entire furca; the duct tapers quickly. 

Autapomorphies.—No character states in 
the current analysis appear to be autapo- 
morphic for this species. Despite this, the 
gold (as opposed to pale) coloration of the 
decumbent pile on the male notum may be 
autapomorphic. 

Distribution.—The known distribution 
for this species is found in Fig. 1. Nearly 
all specimens were collected near 2,000 m 
elevation, with the exceptions of 1 d and 1 


570 


? collected under 500 m in Chamela, Jal- 
isco. The distribution encompasses parts of 
the following biogeographical regions: the 
Provincia Xerdéfila Mexicana, the Provincia 
Mesoamericana de Montafia, and the Prov- 
incia Pacifica. 

Biology.—This species appears to be 
most abundant in the month of July, with a 
period of activity from late June to early 
October. One 6 was collected at a flower 
of Donnellsmithia hintonii Mathias and 
Constance (Asteraceae). 


Ozodiceromyia livdahli Gaimari and 
Irwin, new species 
(Figs. 24, 28) 


Male.—Body length: 7.6 mm. 

Head: 1.11 mm long, 2.12 mm wide, 
1.23 mm high. Distance between eyes at an- 
tennal level 0.60 mm; at genal level 0.98 
mm. Antenna brown, except basal half of 
scape orange. Scape 1.02 mm long, 0.12 
mm wide; with short, fine, black setae, 
evenly distributed over entire scape (includ- 
ing median surface), and few larger setae. 
Pedicel 0.23 mm long, barrel-shaped, setose 
(including median surface). First flagellom- 
ere 0.92 mm long, 0.15 mm wide; with 
short setae covering basal two-thirds. Stylus 
0.11 mm long, inserted subapically. Anten- 
nal base to nearest edge of eye 0.15 mm. 
Frons bulging 0.12 mm beyond eye in lat- 
eral view. Face below antenna with silver 
pruinescence. Parafacial lacking pile, with 
silver pruinescence extending to antennal 
base and dorsally along eye margin halfway 
up frons. Frons otherwise shiny black; with 
few, short, fine, black setae (0.14 mm long) 
dorsolateral to antennal base; remainder of 
frons bare. Genal pile short, darkened. Pal- 
pal pile white basally, dark brown distally. 
Postgenal and occipital pile white; occiput 
with silver pruinescence only along edge of 
eye, and with several black setae. Median 
occipital sclerite flattened; glabrous, shiny 
black; upper edge not rounded. Postocular 
setae black, arranged in single, transverse 
row. Ocellar tubercle with silver pruinesc- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ence, and fine, forward-directed, black se- 
tae. 

Thorax: Scutum and scutellum with ap- 
pressed, flattened, pale [or gold] pile, and 
erect, pale [or gold] pile. Scutum 2.30 mm 
long, 1.80 mm wide; ground color black; 
with silver-grey pruinescence dorsally, 
lacking pruinescence laterally; median vitta 
diffuse bronze; dc vittae faintly present 
only as lines of reduced pruinescence; dc 
setae absent [or | pair present]. Scutellum 
with silver pruinescence, reduced pruinesc- 
ence anteriorly. Halter yellow, except base 
of stalk brown. Katatergite with dense, 
long, white pile. Anepimeron, katepimeron, 
and meron lacking pile, and with reduced 
silver pruinescence, appearing as vertical 
brown stripe from wing base to between 
second and third coxae. Anepisternum with 
silver pruinescence, and with white pile. 
Katepisternum with silver pruinescence, 
with white pile only on vertical crest along 
middle of pleurite. Prepimeron with silver 
pruinescence, lacking pile. 

Legs: Coxae with silver pruinescence, 
posterior surfaces less so. Posterior surface 
of hindcoxa lacking pile. Femora dark 
brown; with appressed, scale-like white and 
brown pile dorsally. Fore- and midfemora 
with erect and long, recumbent, white pile. 
Tibiae with short, erect, black setae; prox- 
imal, dorsal surfaces with sparser setal cov- 
ering, or nearly bare. Foretibia orange ba- 
sally, becoming dark brown distally; clavate 
distally. Midtibia orange, becoming dark- 
ened distally. Hindtibia orange. Foretarsus 
dark brown. Mid- and hindtarsi dark brown 
except basal two tarsomeres mostly orange. 

Wing: 6.18 mm long. Basal costal lobe 
with setae arranged in line along outer 
edge, extending into outer row of costal se- 
tae; second line of setae along base of basal 
costal lobe, extending into inner row of cos- 
tal setae. Entire membrane slightly dark- 
ened; veins and membrane orange basally, 
brown distally. 

Abdomen: Tergites with erect and re- 
cumbent, white pile, except tergite 1 with 
recumbent brown pile medioposteriorly; 


VOLUME 102, NUMBER 3 


dorsally, with silver pruinescence; laterally, 
lacking pruinescence (showing dark brown 
ground color), except tergite 1 and posterior 
edges of basal tergites with silver pruinesc- 
ence. Sternites with silver pruinescence 
only along anterior edge of sternite 2. 

Terminalia (paratype, MEI 038822): 
Sternite 8 with fine black setae restricted to 
posterior edge, which is emarginate medi- 
ally. Tergite 8 dumbell-shaped, with fine 
black setae restricted to posterior edge. 
Epandrium 0.23 mm long, 0.66 mm wide 
at widest point; orange; emarginate anteri- 
orly; dorsal surface with white setae on pos- 
terior half (Fig. 24); lateral edges parallel; 
posterolateral corners extended posteriorly. 
Subepandrial plate attached to posterolat- 
eral and posterior edges; sclerotized portion 
V-shaped posteriorly. Sclerotized portion of 
cerci 0.23 mm long; extending posteriorly 
slightly beyond posterolateral corners of 
epandrium; subequal in length to ventral 
epandrial sclerite. Gonocoxites 0.80 mm 
wide; orange; with white setae (Fig. 28), up 
to 0.45 mm long; fusion 0.36 mm long at 
midline, lacking suture. Inner gonocoxal 
process flanged; knob with several white 
setae. Outer gonocoxal process flange-like, 
0.38 mm long; with small patch of short, 
fine setae dorsodistally. Gonocoxal apode- 
me entirely within anterior edge of gono- 
coxite; lacking sclerotized bridge to para- 
meral sheath of phallus. Ventral gonocoxal 
process 0.23 mm long, tapering distally; 
bare. Ventral lobes distinct, fused basally by 
thin, transparent membrane. Gonostylus ex- 
panded ventrally into large lobe; with dor- 
sobasal lobe; with subapical, lateral spur. 
Dorsal apodemes of aedeagus parallel; sub- 
equal! in length to ventral apodeme; para- 
meral sheath smooth dorsally. Ventral apo- 
deme uniformly wide; lacking ventral keel. 
Ejaculatory apodeme 0.30 mm long; stick- 
like, slightly expanded distally. Lateral 
ejaculatory process a complete ring dorsal- 
ly, but notched; set into aedeagus; lightly 
sclerotized. Distiphallus swollen basally, 
long, recurved; distally parallel with ventral 
apodeme. 


571 


Female.—Similar to ¢ except as follows: 
body length 10.1 mm. 

Head: 1.25 mm long, 2.30 mm wide, 
1.65 mm high. Distance between eyes at 
level of anterior ocellus 0.51 mm; at anten- 
nal level 0.96 mm; at genal level 1.04 mm. 
Scape 1.29 mm long, 0.17 mm wide. First 
flagellomere 1.04 mm long, 0.17 mm wide. 
Antennal base to nearest edge of eye 0.30 
mm. Frons bulging 0.15 mm beyond eye in 
lateral view. Frons shiny black; lower frons 
with short (up to 0.09 mm long) setae dor- 
solateral to antennal base; upper frons dis- 
tinctly bulging and distinct from lower 
frons, with small patch of setae (up to 0.09 
mm long) between edge of eye and midline. 
Median occipital sclerite and upper edge 
rounded; transverse row of black setae 
across median occipital sclerite in addition 
to row of postocular setae. 

Thorax: Scutum and scutellum with ap- 
pressed and semi-appressed white pile. Scu- 
tum 2.75 mm long, 1.95 mm wide. Pile of 
anepisternum and katepisternum short, 
white. 

Wing: 7.26 mm long. 

Abdomen: Tergites and sternites mostly 
lacking pruinescence; tergites with short, 
fine appressed brown and white pile. 

Terminalia: Furca 0.54 mm long, 0.32 
mm wide; anterior edge not sclerotized; an- 
terolateral prongs dorsoventrally flattened, 
separation subequal to greatest width of fur- 
ca. Furcal bulla not sclerotized; posteriorly 
with indented cavity. Gonopore basal to 
furcal bulla, originating within posterior 
cavity. Common spermathecal duct basally 
0.11 mm wide, tapering to 0.05 mm within 
0.33 mm distance from gonopore; 0.93 mm 
long. Spermathecal ducts originate from 
clean trifurcation with central sac duct; cen- 
tral sac duct wider in basal diameter than 
spermathecal ducts. Central sac duct 0.45 
mm long. Central sac 0.68 mm long, 0.39 
mm wide. Spermatheca 0.18 mm in diam- 
eter; rounded, but basal edge slightly flat- 
tened. 

Type materials—HOLOTYPE 3d (MEI 
038101) with the following labels: 5 mi. E 


572 


Fort Apache, ARIZ[ona]., VII-28—1964, E. 
I. Schlinger. This pinned specimen is in ex- 
cellent condition, and is deposited in 
CASC. 

Materials examined.—PARATYPES. 
MEXICO: Chihuahua, D. J. Knull, J. N. 
Knull, 22-VII-1961 (1 2, MEI 038853, 
SDGC), 29-VII-1955 (1 2, MEI 038858, 
SDGC) ; J. N. Knull, 18-VIII-1936 (1 4, 
MEI 038854, OSUC); 2,072.64 m, J. A. 
Chemsak, 14-VII-1964 (2 36, MEI 038817, 
038818, EMEC), J. A. Chemsak, J. A. Pow- 
ell, 14-VII-1964, black and white lights (1 
36, MEI 038837, EMEC), J. A. Powell, 15- 
VII-1964 (1 63, MEI 038815, EMEC); 
2,011.68 m, D. Rockefeller, Gertsch Exp., 
21-VII-1947 (1 2, MEI 038791, AMNH); 
8.1 km W of Parrita, Santa Clara Canyon, 
D. D. Linsdale, 3-IX-1956 (1 9, MEI 
052154, USNM), J. W. MacSwain, 3-IX- 
1956 (1 6, MEI 052159, USNM; | 3, MEI 
038874, 2 2, MEI 038873, 038875, EMEC; 
1 2, MEI 052158, SDGC); 20.9 km E of 
Cuauhtemoe, 2,011.68 m, J. A. Chemsak, 
11-VI-1964 (1 2, MEI 038780, EMEC); 
La Bufa, Sierra Madre Mountains, 900 m, 
D. D. Giuliani, 7-VII-1972 (1 6, MEI 
052246, SDGC); Sinaloa, bet.[ween] NW 
of Culiacan & Las Mochis, D. Spencer, R. 
Ryckman, J. Ryckman, A. Ryckman, 21- 
VII-1957 (1 6, MEI 038787, UNAM); ca. 
100 km NW of Culiacan, D. Spencer, R. 
Ryckman, J. Ryckman, A. Ryckman, 21- 
VITI-1957 (1 36, MEI 038779, UNAM). 

UNITED STATES: Arizona, Cochise 
County, L. D. Anderson, 21-VIII-1966, at 
light (1 2, MEI 040761, SDGC) ; M. Sta- 
tham, 19-VIII-1959 (1 2, MEI 053688, 
AMNH); Mina Canyon, 20-VII-1925 (1 6, 
MEI 041774, DENH); Chiricahua Mts., J. 
K. Robertson, 1-VIII-1965 to 2-VIII-1965 
(2 2, MEI 030930, 030931, LACM); K. W. 
Brown, 7-VIII-1965, UV light (1 36, MEI 
038868, INBC; 1 2, MEI 038865, ZMAS); 
R. M. Bohart, 6-VIII-1958 (1 3, MEI 
038915, NHMW); 1.6 km S of Portal, E. 
G. Linsley, J. M. Linsley, 16-VITI-1966 (1 
2, MEI 038870, EMEC) ; 1.6 km SW of 
Portal, J. A. Powell, 11-VII-1972 to 18-VII- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


1972 (1 6, MEI 038825, ZMAS; 1 36, MEI 
038826, NMSA; 1 36, MEI 038827, INHS; 
Le 3) MERns038828;, ASUT, 41isd.eMeI 
038829, BMNH; 1 6, MEI 038830, 
MEIC); 1.61 km E, 8.05 km N of Portal, 
2-IX-1959, at light (1 6, MEI 043432, 
UMSP) ; 2.7 km W of Portal, S. I. From- 
mer, S. L. Frommer, 22-VIII-1975, dry 
stream bed (1 6, MEI 052194, UCRC); 3.2 
km NE Portal, 1645.92 m, J. Wilcox, 20- 
IX-1962 (1 3, MEI 038878, MEIC); 4.8 
km S of Paradise, A. B. Patterson, 9-VIII- 
1966, malaise trap (1 6, MEI 038813, 
SDGC); 4.8 km W of Portal, D. P. Levin, 
28-VIII-1971, black light (1 2, MEI 
034099, DENH); 8.1 km W of Portal, C. G. 
Moore, 8-VIII-1958 (1 2, MEI 038912, 
MNHN), G. B. Pitman, 19-VIII-1958 (1 6, 
MEI 038904, IZAS; 1 3, MEI 038916, 
BYUC), P. M. Marsh, 13-VIII-1958 (1 6, 
MEI 029552, SDGC), P. Opler, 11-VIII- 
1958 (1 2, MEI 056915, EMEC), 1,645.92 
m, V. D. Roth, 9-VIII-1965 (1 3d, MEI 
038903, UNAM); 9.7 km N of Portal, L. D. 
Anderson, M. D. Anderson, 20-VIII-1970 
(1 2, MEI 038848, SDGC); Cave Creek 
Canyon, 5,000’, D. R. Corr, 7-VIII-1986, 
malaise trap (1 6, MEI 110058, MTEC), 
6.4 km SSW of Portal, R. Davidson, 24- 
VIII-1981, malaise trap (1 6, MEI 052169, 
CMNH); Cave Creek Ranch, G. R. Ballmer, 
15-VIII-1965, UV light (1 2, MEI 038861, 
FMNH), R. Silberglied, 16-VIII-1966, 
black light (1 2, MEI 038866, CUIC), 1524 
m, K. W. Brown, 10-VIII-1965 (1 2°, MEI 
038863, ESUW), M. E. Irwin, 13-VIII- 
1965 (1 2, MEI 038893, MHNG), Portal, 
E. G. Linsley, 1-VIII-1972 to 3-VIII-1972 
(1 2, MEI 038809, EMEC), J. A. Powell, 
2-VII-1972 to 4-VII-1972 (1 6, MEI 
038831, BPBM; 2 36, MEI 038832, 038833, 
2 2, MEI 038834, 038835, EMEC; 1 &, 
MEI 038836, ASUT), 3-VII-1972 to 4-VIH- 
1972 (1 36, MEI 038822, MEIC; 1 3, MEI 
038823, CSUC; 1 6, MEI 038824, TAMU; 
1 3, MEI 038821, 2 2, MEI 029554, 
038820, EMEC; 1 2, MEI 038819, 
MUSM), R. Silberglied, 21-VIII-1966, 
black light—UV (1 6, MEI 038869, 


VOLUME 102, NUMBER 3 


CUIC), 1.6 km S of Portal, 21-VIII-1969 to 
26-VIII-1969 (1 3, MEI 038843, NMSU), 
E. G. Linsley, J. M. Linsley, 15-VIII-1970 
to 20-VIII-1970 (1 6, MEI 038844, EBCC; 
1 36, MEI 038842, 1 2, MEI 038845, 
MEIC; 1 2, MEI 038846, CICESE; 1 °, 
MEI 038847, EMEC; 1 ¢, MEI 038839, 
UCMC; | 2, MEI 038840, IZAV; 1 2°, MEI 
038841, MZSP); Chiricahua National Mon- 
ument, L. M. Martin, 23-VIII-1951 (1 @, 
MEI 030937, LACM); Paradise, G. R. Ball- 
mer, 6-VIII-1966, malaise trap (1 3d, MEI 
038889, ZMHB; 1 3, MEI 038890, MZSP; 
1 3, MEI 038891, MRSN; 1 &, MEI 
038909, CASC; 1 2, MEI 038910, TAMU; 
1 2, MEI 038911, UNAM); Portal, E. I. 
Schlinger, 11-VIII-1967 (1 2, MEI 038849, 
SDGC; 1 2, MEI 038850, MEIC), J. A. 
Powell, 2-VII-1972 to 4-VH-1972, at light 
(1 36, MEI 038792, EMEC), R. M. Bohart, 
15-VIII-1958 (1 2, MEI 029555, SDGC), 
1524 m, H. E. Evans, 9-VIII-1959 (1 9, 
MEI 038812, ZMUC); Rucker Canyon, R. 
M. Bohart, 24-VIII-1979 (1 6, MEI 
052226, UCDC); Southwestern Research 
Station, 8.1 km W of Portal, P. D. Hurd, 3- 
VIII-1958 (1 2, MEI 038872, NMSU), 11- 
VIII-1958 (1 2, MEI 038871, EMEC), 
South Fork Cave Creek, P. H. Arnaud, 5- 
IX-1959 (1 3, MEI 052249, 1 2, MEI 
052248, CASC), 1645.92 m, 14-VIII-1970 
(1 2, MEI 038838, MEIC), H. E. Evans, 
18-VIII-1959 (1 2, MEI 038811, UNAM), 
M. Statham, 26-VII-1957 (1 6, MEI 
052232, AMNH), 27-VII-1959 (1 2, MEI 
052233, AMNH), M. S. Wasbauer, 8-VIII- 
1978 to 9-VIII-1978, malaise trap (1 ¢, 
MEI 071419, CNCI; 1 6, MEI 071420, 1 
2, MEI 071440, CDFA), 10-VIII-1978 to 
11-VIII-1978, malaise trap (1 2, MEI 
071426, CDFA), 14-VIII-1978 to 15-VIII- 
1978, malaise trap (1 6, MEI 071421, 
WSUG; 1) 6.) MEI071422, SDGE;.2°-2, 
MEI 071424, 071425, CDFA), 16-VIII- 
1978 to 17-VIII-1978, malaise trap (1 <4, 
MEI 071448, CDFA), V. D. Roth, 9-IX- 
1979 (2 2, MEI 026953, 026954, SWRS), 
15-VIII-1965 to 20-VIII-1965 (1 6, MEI 
026986, SWRS); Sulphur Draw, G. R. Ball- 


573 


mer, 7-VIII-1965, UV light (1 3, MEI 
038862, NVDA); Sunny Flat, 6.4 km W of 
Portal, C. W. Melton, 27-VIII-1979, black 
light (1 2, MEI 071409, CDFA; 1 2, MEI 
070409, CSUC); Tex Canyon, E G. An- 
drews, 6-VIII-1967 (1 6, MEI 038851, 
DEIC); Dragoon Mountains, Cochise 
Stronghold, R. J. Shaw, 12-VIII-1970 to 
16-VIII-1970, UV light (1 2, MEI 076933, 
ANIC); Huachuca Mountains, D. J. Knull, 
J. N. Knull, 20-VII-1937 (1 92, MEI 
052160, USNM), J. S. Hine, 28-VII-1907 
(1 2, MEI 052222, OSUC), R. L. Westcott, 
16-VIII-1966 (1 2, MEI 079534, WFBM); 
13 km SE of Sunnyside, 1,813.56 m, R. R. 
Snelling, 23-VII-1971 (1 3d, MEI 030927, 
1 2, MEI 030932, LACM); Ash Canyon, 
N. McFarland, 15-[X-1983 (1 6, MEI 
OS521615 1 2eIME!, 0521622 3USNM); 
1554.48 m, N. McFarland, 13-VIII-1982, 
UV light (1 2, MEI 052170, SDMC), 
1676.4 m, R. R. Snelling, 20-VIII-1971 (2 
3, MEI 030928, 030940, LACM); Carr 
Canyon, E. P. Van Duzee, 5-VIII-1924 (1 
6, MEI 029553, MCZC), J. O. Martin, 6- 
VIII-1924 (1 2, MEI 038816, MCZC), 
1,645.92 m, C. W. O’Brien, 7-IX-1995, 
black light (1 ¢, MEI 038782, EMEC), H. 
B. Leech, J. W. Green, 8-VIII-1952 to 9- 
VITI-1952, floor of Carr Canyon (1 6, MEI 
052234, CASC); Copper Canyon, W. E 
Barr, 5-VIII-1990, beat. Quercus (1 36, MEI 
079905, 1 2, MEI 079904, WFBM), at Rd. 
61.) Sec. 10, F238) /R29E, W: Fy Bars, 31- 
VII-1979 (1 2, MEI 079468, NYSM), 
1,828.8 m, Y. F Hsu, J. Powell, M. Prentice, 
3-VIII-1989, black light—UV (1 3, MEI 
081462, EMEC); Miller Canyon, G. E. 
Wallace, 24-VIII-1965, UV light (1 ¢, MEI 
038855, NYSM; 1 &, MEI 038852, 
OSUC); Ramsey Canyon, L. D. Anderson, 
21-VIII-1969 (1 2, MEI 038888, ANSP); 
Stump Canyon, Olson, 31-VII-1979, UV 
light (1 36, MEI 076939, UAIC); Gila 
County, Pinal Mountains, Sixshooter Can- 
yon, 1,524 m (1 2, MEI 052156, USNM); 
Graham County, Graham Mountain, Noon 
Creek, E G. Werner, 28-VII-1954, at light 
(1 3, MEI 038905, MEIC), G. D. Butler, 


574 


1-VIII-1957 (1 2, MEI 038913, SDGC); 
Noon Creek Camp, L. G. Bezark, G. M. 
Nishida, C. Kitayama, B. Tilden, 29-VIII- 
1875 30N fhisht (lid, MEL. 041767, 
UCMC), Highway 266, L. G. Bezark, G. 
M. Nishida, C. Kitayama, B. Tilden, 29- 
VIII-1975 to 30-VIII-1975, UV light (1 @, 
MEI 041771, MEIC; 1 ¢&, MEI 041772, 
NVDA); Pinalefo Mts., Hospital Flats, Mt. 
Graham, G. D. Butler, 15-VIII-1953 (1 &, 
MEI 076943, EMEC; 1 2, MEI 076928, 
KSUC); Maricopa County, 3.22 km W of 
Tortilla Flat, 505.968 m, Canyon Lake, J. 
LaSalle, S. Y. H. Lin, 22-VIII-1982, black 
light (1 2, MEI 043088, UCRC) ; Pima 
County, Baboquivari Mountains, Brown 
Canyon, E G. Werner, W. Nutting, 4-VIII- 
1961, UV light (1 d, MEI 038884, UAIC; 
1S IMEI 038877, 0S DGC." ti -oe MET 
038879, DEIC); Santa Catalina Mountains, 
S. L. Wood, J. B. Karen, 9-VIII-1962, black 
light (1 2, MEI 037668, BYUC), Bear 
Canyon, 19.32 km HK Highway, FE G. Wer- 
ner, W. Nutting, 26-VII-1961, light trap (1 
2, MEI 038899, UAIC) , Molino Basin, 
1,402.08 m, C. W. O’Brien, L. B. O’Brien, 
4-IX-1965, black light (1 2, MEI 038814, 
SDGC), Sabino Basin, C. H. Townsend, 20- 
VIII (3, MET 052157, USNM); Santa 
Rita Mountains, Box Canyon, R. EK Denno, 
16-VIII-1970 (1 2, MEI 052224, UCDO), 
Madera Canyon, L. G. Bezark, G. M. Nish- 
ida, C. Kitayama, B. Tilden, 31-VIII-1975 
(1 6, MEI 041769, ESUW), W. J. Hanson, 
4-IX-1968 (1 6, MEI 070469, EMUS; | 6, 
MEI 071469, CDFA), 1569.72 m, J. M. 
Sheppard, 8-VIII-1969, black light, oaks (1 
2, MEI 052193, OSEC), N End Rosemont 
Area, 31°48-53'N, 110°42—47'W, 4,400-— 
6,175’ El (6,000’), Anamax Mine Inventory 
1975-1976, Wasp Cn. Sec. 31, J. Busacca 
and C. Olson, 8-28-1975, U V light (1 &, 
MEI 076921, UAIC); Santa Rita Res. 
Range, D. K. Faulkner, 5-VIII-1980 (1 6, 
MEI 052164, 1 2, MEI 052166, SDMC); 
Pima and Santa Cruz Counties, Santa Rita 
Mountains, E. L. Todd, 1-VII-1941 (1 @, 
MEI 052221, SEMC), FE H. Parker, 12-VIII- 
1935 (1 ¢, MEI 052155, USNM), S. L. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Szerlip, J. A. Powell, 13-VIII-1974, at light 
(1 36, MEI 052175, KSUC); Santa Cruz 
County, K. Roever, 3-VIII-1959, black light 
trap (1 2, MEI 038886, UAIC; 1 2, MEI 
038887, IZAS); Canelo, G. D. Butler, 3- 
VIII-1965 (1 2, MEI 038901, BPBM); Ca- 
nelo Hills, Parker Cyn, R. L. Westcott, 30- 
VII-1979 (1 3, MEI 079463, CNCI); Pena 
Blanca, Werner, Olson, 11-VIII-1983 (1 2, 
MEI 076929, UAIC); Sycamore Canyon, 
near Ruby, K. Roever, 2-VIII-1959, light 
trap (1 2, MEI 038876, UNAM; 1 2°, MEI 
038885, INBC; 1 2, MEI 038900, UNSM) 
V. L. Versterby, 6-IX-1963 (1 @, MEI 
038880, MEIC); Patagonia Mountains, F 
G. Werner, G. D. Butler, 8- VITI-1955, west- 
ern slope (1 6, MEI 038867, SDGC), 9- 
VIII-1966, western slope (1 2, MEI 
038918, MEIC), G. D. Butler, E G. Werner, 
9-VIII-1955, western slope (1 ¢6, MEI 
038906, QCAZ; 1 3, MEI 038907, ANSP; 
1 3, MEI 038908, SDGC; 1 6, MEI 
038914, UAIC; 1 36, MEI 038917, UNSM), 
Mount Washington, 1676.4 m, L. G. Be- 
zark, R. A. Cunningham, D. E. Russell, 12- 
VIII-1991 to 13-VIII-1991, Hg vapor and 
UV blacklight (1 2, MEI 038115, WSUC); 
Pajarito Mountains, R. H. Arnett Jr, E. 
VanTassell, 6-VIII-1961 (1 2, MEI 038784, 
FSCA); Santa Rita Mountains, Madera 
Canyon, 3-VIII-1977 (1 36, MEI 052167, 
SDGC), A. J. Gilbert, N. J. Smith, 11-VIII- 
1981 (1 2, MEI 038119, SDGC; 1 2, MEI 
038120, MEUC), D. K. Faulkner, 9-VIII- 
1978 to 20-VIII-1978 (1 2, MEI 052168, 
ZMAS), D. J. Knull, J. N. Knull, 1-VIII- 
1972 (2 Y, MEI 052223, TAUD, Da 
Faulkner, 20-VIII-1979 (1 2, MEI 052165, 
BMNH), E. M. Fisher, 7-VIII-1962 to 9- 
VIII-1962 (1 2, MEI 030943, CMNH; 1 °, 
MEI 030929, CNCI; | 2, MEI 031008, 
EMUS; 1 6, MEI 030942, KUIC), G. R. 
Ballmer, K. Brown, 31-VII-1965 (1 3, MEI 
038860, 1 2, MEI 038859, MEIC; 1 °, 
MEI 038808, NHRM), L. G. Bezark, G. M. 
Nishida, C. Kitayama, B. Tilden, 24-VIII- 
1975, UV light (1 6, MEI 041768, NHRM; 
1 3, MEI 041773, UMRM; 1 2, MEI 
041770, MEIC), L. M. Martin, 14-VIII- 


VOLUME 102, NUMBER 3 


1949 (1 2, MEI 030936, MRSN), 15-VIII- 
1949 (1 36, MEI 030934, LACM; 1 2, MEI 
030933, CNCI; 1 2, MEI 030935, UMSP), 
16-VIII-1949 (1 3, MEI 030941, LACM), 
31-VII-1947 (1 2, MEI 030939, INHS), M. 
A. Cazier, R. Schrammel, C. Vaurie, P. Vau- 
ne. 13-Vil-1952.(1. 2 MEP, 038781; 
AMNB), P. H. Sullivan, 31-VIII-1970 (1 @, 
MEI 038856, KUIC), R. L. Westcott, 13- 
VII-1964 (1 6, MEI 079570, WFBM; 1 °, 
MEI 079569, FSCA), W. A. McDonald, 17- 
VIII-1955 to 20-VIII-1955 (1 ¢, MEI 
038902, LACM), 1,402.08 to 1,706.88 m, 
4-VIII-1975 (1 3, MEI 076916, CICESE; 
1 3, MEI 076918, FMNH; 2 6, MEI 
076919, 076915, 1 2, MEI 076920, UAIC; 
1, 22; MEL 076940, ‘-ZMHB:.«1. 2... MEI 
O76917. -EBCE: a1. SieMEL 076914, 
NHMW), 1,487.424 m, J. G. Franclemont, 
20-VII-1959 (1 °°, MEI 038810, CUIC), L. 
M. Martin, 20-VIII-1946 to 29-VIII-1946 
(1 2, MEI 031062, EMEC; 1 2, MEI 
030938, CNCD, V. L. Versterby, 8-IX-1963 
(1 2, MEI 038894, QCAZ; 1 2, MEI 
038895, MEIC), 12-VIII-1963 (1 6, MEI 
038896, MUSM; | 36, MEI 038897, ANIC; 
1 6, MEI 038898, IZAV), 21-VII-1963 (1 
36, MEI 038881, MHNG), 22-IX-1963 (1 
?, MEI 038883, UMRM), 31-VII-1963 (1 
36, MEI 038882, MEUC), 1,524 m, K. W. 
Brown, P. Petrulis, 3-[X-1971, UV light (1 
6, MEI 033978, 1 @%, MEI 033979, 
PMNH), 1,554.48 m, D. R. Davis, 10-VII- 
1964 to 26-VII-1964 (1 3, MEI 052152, 
SEMC; 1 2, MEI 052153, NMSA), Bog 
Springs, 1,706.88 m, E. Lindquist, 6-VIII- 
1973 (1 36, MEI 052231, CNCI); County 
unknown, Floricera, Sta Rita Mts., Olson, 
Burme, Frank, 30-VII-1980 (1 ¢, MEI 
076938, UAIC); Garces, N. Banks, VIII (1 
2, MEI 038864, MCZC) ; New Mexico, Hi- 
dalgo County, 1,645.92 m, Skeleton Can- 
yon, G. R. Ballmer, 12-VIII-1965 (1 ¢, 
MEI 038892, USNM); 56.35 km E of 
Douglas, Arizona, C. W. Sabrosky, 22-IX- 
1965 (1 2, MEI 038857, USNM); Coro- 
nado National Forest, Peloncillo Moun- 
tains, 1.4 km W National Forest Boundary, 
Black C.C.C. Dam Road, NFR63, 1661.16 


S/S 


m, M. A. Metz, 28-VI-1997, malaise trap 
(2 36, MEI 103430, 103426, SDGC; 2 6, 
MEI 103431, 103432, MEIC; 1 6, MEI 
103433, OSEC; 1 3d, MEI 103427, ZMUC), 
4.0 km W National Forest Boundary Black 
C.C.C. Dam Road, NFR63, 1,682.496 m, 
M. A. Metz, 28-VI-1997, malaise, 9 m 
Fock’s (1 ¢, MEI 103428, INHS; 1 6, MEI 
103429, ZMAS; 1 6, MEI 103424, TAUI; 
1 3d, MEI 103425, MNHN). 

Diagnosis.—The scape in this species is 
setose, but most of the setae are short, with 
few longer ones. The lower frons bulges 
only slightly, with silver pruinescence lat- 
erally, extending down along parafacial. 
The face below the antenna also has silver 
pruinescence. In males, the notum is dense- 
ly covered with erect and appressed pale or 
gold pile. In females, the notum and ab- 
dominal tergites are covered with short, 
fine, appressed brown and white pile. In the 
male genitalia, the setae of the epandrium 
and gonocoxites are orange. The subepan- 
drial plate is attached to the epandrium at 
the posterolateral corners, and the sclero- 
tized portion of the subepandrial plate is V- 
shaped posteriorly. The gonocoxites lack a 
suture along the midline separating the two 
lateral halves. The outer gonocoxal process 
is widened distally, with a small patch of 
setae on the dorsolateral surface. The ven- 
tral gonocoxal process of the gonocoxites is 
present, elongated and flattened; there are 
no clumped setae at its base. The base of 
the ventral lobes is U-shaped and broad be- 
tween the two halves. The parameral sheath 
of the aedeagus is smooth dorsally. The 
ventral apodeme lacks a ventral keel. The 
distiphallus is swollen basally, and is re- 
curved and parallel with the ventral apo- 
deme at the tip. In the female terminalia, 
the furca is not notched posteriorly, and 
lacks a posterolateral peg. The anterolateral 
furcal prongs are separated by a distance 
subequal to the greatest width of the furca. 
The common spermathecal duct is broad 
basally, at half the width of the entire furca; 
the duct tapers quickly. 


Rip: 

Autapomorphy.—Gonocoxal setae pale 
(character 26, state 0). 

Distribution.—The known distribution 
for this species is found in Fig. 2. Most of 
the localities fall within an elevational 
range of 1,000—2,000 m, although 1 2 was 
collected near 500 m. The distribution en- 
compasses parts of the following biogeo- 
graphical regions: the Provincia Xer6fila 
Mexicana, the northern part of the Provin- 
cia Mesoamericana de Montafia, and the 
montane regions of the southeastern quarter 
of Arizona and southwestern New Mexico. 

Etymology.—Patronym for Todd P. Li- 
vdahl, professor of Biology at Clark Uni- 
versity, Worcester, Massachusetts, alma 
mater of SDG; to be treated as a noun in 
apposition. 

Biology.—This species is most active in 
July and August, with a range from the end 
of June (4 d<d) through September. This 
species seems to be commonly collected at 
UV and other light sources at night. 


Ozodiceromyia mexicana Bigot 
(Figs. 9-10, 16, 21, 25, 30-31, 34-35, 
41) 

Ozodiceromyia mexicana Bigot 1890: 321. 

Type locality: Mexico. Type: HT? in 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Bae 


Known distribution for Ozodiceromyia livdahli. 


BMNH. Bigot 1890: 323 (key ref.); 
Krober 1912: 211 (repr. orig. descr.), 
1913: 8 (cat. cit.); Cole 1923: 19—20 (an- 
not. listing); Gaimari 1998: 124-125 (lit. 
list), 157-159 (in phylogeny). 

Ozodiceromya (sic) mexicana: Bigot 1890: 
321 (orig. descr.); Irwin and Lyneborg 
198la: 196 (illust.: coxa), 199 (illust.: an- 
tenna), 254 (illust.: ¢ genit.), 257 (list- 
ing). 

Ozodiceronyma (sic) mexicana: Godman 
1901: 378 (listing). 

Euphycus setosus Kréber 1912: 211. Type 
locality: Mexico. Type: LTd (designated 
herein) in NHMW. Irwin and Lyneborg 
1981a: 257 (prop. syn.). 

Ozodiceromya (sic) setosa: Irwin and Ly- 
neborg 198la: 257 (comb. change). 


Female holotype.—The HT?® is in poor 
condition, i.e., the abdomen and thorax are 
completely covered with orange fungus. 
Fortunately, the head is mounted separately 
and is fungus-free. The head is distinct 
enough to associate this specimen with a 
recognizable species. The following de- 
scription deals only with the characteristics 
that can be seen in the specimen. Following 
this, the full redescription of the recognized 


VOLUME 102, NUMBER 3 


species comes from the LT¢ of the junior 
synonym, Euphycus setosus Krober. 

Body length: 7.1 mm. 

Head: 1.13 mm long, 1.73 mm wide, 
1.28 mm high. Distance between eyes at 
level of anterior ocellus 0.33 mm; at anten- 
nal level 1.01 mm; at genal level 0.96 mm. 
Scape 0.59 mm long, 0.12 mm wide; with 
long, fine and thick, black setae, evenly dis- 
tributed over entire scape (including median 
surface). Pedicel 0.18 mm long, barrel- 
shaped, setose (including median surface). 
Antennal base to nearest edge of eye 0.20 
mm. Frons bulging 0.17 mm beyond eye in 
lateral view. Parafacial lacking pile; silver 
pruinescence in thin line along edge of eye; 
face and parafacial otherwise glabrous, 
shiny black. Frons shiny black; lower frons 
globose and bulging to antennal insertion; 
upper frons distinct from lower frons, bulg- 
ing. Genal pile short and black. 

Legs: Foreleg dark brown, except fore- 
tibia paler and lacking setae on proximal, 
dorsal surface. Midleg dark brown, except 
midtibia orange basally. 

Wing: 5.85 mm long. 

Male.—Lectotype of Euphycus setosus 
Kroéber, to represent the species Ozodicer- 
omyia mexicana. 

Body length: 6.9 mm. 

Head (Fig. 9): 0.90 mm long, 1.59 mm 
wide, 1.35 mm high. Distance between eyes 
at antennal level 0.83 mm; at genal level 
1.01 mm. Antenna dark brown. Scape 0.71 
mm long, 0.18 mm wide; with long, fine 
and thick, black setae, evenly distributed 
over entire scape (including median sur- 
face). Pedicel 0.20 mm long, barrel-shaped, 
setose (including median surface). First fla- 
gellomere 0.60 mm long, 0.15 mm wide; 
with short setae covering basal two-thirds. 
Stylus 0.09 mm long, inserted subapically. 
Antennal base to nearest edge of eye 0.26 
mm. Frons globose, bulging 0.18 mm be- 
yond eye in lateral view. Parafacial lacking 
pile, with silver pruinescence only along 
eye edge; pruinescence extends dorsally in 
thin line along eye margin halfway up 
frons; face and parafacial otherwise gla- 


577 


brous, shiny black. Frons otherwise shiny 
black; with long, fine, black setae (up to 
0.38 mm long); upper frons bare. Genal pile 
short, darkened. Palpal pile white basally, 
dark brown distally. Postgenal pile white; 
occipital pile black; occiput with silver 
pruinescence only along edge of eye, and 
with several black setae. Median occipital 
sclerite flattened; glabrous, shiny black; up- 
per edge not rounded. Postocular setae 
black, arranged in single, transverse row. 
Ocellar tubercle shiny black, with fine, for- 
ward-directed, black setae. 

Thorax: Scutum and scutellum with 
erect black pile sparsely interspersed with 
appressed gold pile. Scutum 2.08 mm long, 
1.70 mm wide; ground color black; with sil- 
ver-grey pruinescence dorsally, lacking 
pruinescence laterally; median vitta diffuse 
bronze; dc vittae absent; dc setae absent [or 
1 pair present]. Scutellum with silver prui- 
nescence, reduced pruinescence anteriorly. 
Halter yellow. Katatergite with dense, long, 
orange pile. Anepimeron, katepimeron, and 
meron lacking pile, and with reduced silver 
pruinescence, appearing as vertical brown 
stripe from wing base to between second 
and third coxae. Anepisternum with silver 
pruinescence, and with white pile. Katepis- 
ternum with silver pruinescence, with white 
pile only on vertical crest along middle of 
pleurite. Prepimeron with silver pruinesc- 
ence, lacking pile. 

Legs: Coxae with silver pruinescence, 
posterior surfaces less so. Posterior surface 
of hindcoxa with long, white pile (Fig. 16). 
Femora dark brown. Fore- and midfemora 
with appressed, scale-like, white pile, and 
with long, erect, recumbent, white and 
black pile. Hindfemur with appressed, 
scale-like brown and white pile. Tibiae with 
short, erect, black setae; proximal, dorsal 
surfaces with sparser setal covering, or 
nearly bare. Foretibia orange basally, be- 
coming dark brown distally; clavate distal- 
ly. Midtibia orange, becoming darkened at 
distal tip. Hindtibia orange basally, becom- 
ing dark brown distally. Foretarsus dark 
brown. Mid- and hindtarsi dark brown ex- 


578 


cept for basal two tarsomeres orange at 
their bases. 

Wing: 6.24 mm long. Basal costal lobe 
with setae arranged in line along outer 
edge, extending into outer row of costal se- 
tae; second line of setae along base of basal 
costal lobe, extending into inner row of cos- 
tal setae. Entire membrane slightly dark- 
ened; veins and membrane orange basally, 
brown distally. 

Abdomen: Tergite 1 with erect and re- 
cumbent gold pile, except dark brown me- 
dioposteriorly; remaining tergites with erect 
and recumbent, white pile; dorsally, with 
silver pruinescence; laterally, lacking prui- 
nescence (showing dark brown ground col- 
or), except tergite 1 and posterior edge of 
tergite 2 with silver pruinescence. Sternites 
with silver pruinescence only along anterior 
edge of sternite 2. 

Terminalia (MEI 052240): Sternite 8 
with black setae along posterior edge, 
which is emarginate medially. Tergite 8 
dumbell-shaped, with black setae along 
posterior edge. Epandrium (Fig. 25) 0.24 
mm long, 0.63 mm wide at widest point; 
orange; emarginate anteriorly; dorsal sur- 
face with white setae on posterior two- 
thirds; lateral edges parallel; posterolateral 
corners extended posteriorly. Subepandrial 
plate (Fig. 25) unattached to epandrium; 
sclerotized portion reduced, divided into 
two small plates. Sclerotized portion of cer- 
ci 0.23 mm long; extending posteriorly be- 
yond posterolateral corners of epandrium; 
subequal in length to ventral epandrial 
sclerite. Gonocoxites (Figs. 30-31) 0.77 
mm wide; orange; with white setae, up to 
0.50 mm long; fusion 0.35 mm long at mid- 
line, with suture. Inner gonocoxal process 
flanged, knob with several white setae. Out- 
er gonocoxal process flange-like, 0.23 mm 
long; bare. Gonocoxal apodeme entirely 
within anterior edge of gonocoxite; lacking 
sclerotized bridge to parameral sheath of 
phallus. Ventral gonocoxal process absent, 
lateral part of corresponding edge with 
dense patch of fine orange setae, up to 0.30 
mm long. Ventral lobes distinct, fused ba- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


sally by thin, transparent membrane. Gon- 
ostylus expanded ventrally into large lobe; 
with dorsobasal lobe; lacking subapical, lat- 
eral spur. Dorsal apodemes (Fig. 34) of ae- 
deagus parallel; subequal in length to ven- 
tral apodeme; parameral sheath with dis- 
tinct transverse wrinkles dorsally. Ventral 
apodeme thin, of uniform width; lacking 
ventral keel. Ejaculatory apodeme 0.35 mm 
long; stick-like, slightly expanded distally. 
Lateral ejaculatory process a complete ring 
dorsally, but notched; set into aedeagus; 
lightly sclerotized. Distiphallus long, re- 
curved; distally parallel with ventral apo- 
deme (Fig. 35). 

Female (MEI 052163).—Similar to 6 
except as follows: body length 8.6 mm. 

Head (Fig. 10): 1.10 mm long, 1.76 mm 
wide, 1.28 mm high. Distance between eyes 
at level of anterior ocellus 0.32 mm; at an- 
tennal level 0.83 mm; at genal level 0.95 
mm. Scape 0.81 mm long, 0.15 mm wide. 
First flagellomere 0.71 mm long, 0.15 mm 
wide. Antennal base to nearest edge of eye 
0.32 mm. Frons shiny black; lower frons 
globose and bulging to antennal insertion, 
with fine, black setae (up to 0.12 mm long) 
in patch above antennal base; upper frons 
bulging and distinct from lower frons. 

Thorax: Scutum and scutellum with re- 
cumbent black pile and appressed gold pile. 
Scutum 2.08 mm long, 1.80 mm wide. Ane- 
pisternum with reduced silver pruinescence; 
pile of anepisternum and katepisternum 
sparse, short, white. 

Wing: 6.12 mm long. 

Abdomen: Tergites and sternites lacking 
pruinescence (showing dark brown ground 
color); tergites with appressed and semi-ap- 
pressed gold pile. 

Terminalia (paratype, MEI 070426) (Fig. 
41): Furca 0.53 mm long, 0.38 mm wide; 
anterior edge not sclerotized; anterolateral 
prongs dorsoventrally flattened, separation 
subequal to greatest width of furca; pos- 
terolaterally with small, ventromedially ori- 
ented peg; posterior edge notched. Furcal 
bulla not sclerotized; posteriorly with in- 
dented cavity. Gonopore basal to furcal bul- 


VOLUME 102, NUMBER 3 


la, originating within posterior cavity. Com- 
mon spermathecal duct 1.14 mm long. 
Spermathecal ducts originate from clean tri- 
furcation at base of central sac duct; central 
sac duct absent. Central sac 0.89 mm long, 
0.30 mm wide. Spermatheca 0.15 mm long, 
0.15 mm wide; rounded distally, but basal 
edge flattened. 

Type materials—HOLOTYPE 2°: (MEI 
103180) top label: Mexic[o]; with the fol- 
lowing handwritten, second label (note, this 
is a standard provisional label of Bigot 
which he never updated): n. genus, Ozodi- 
ceromyia, J. Bigot, O. mexicana 2, n. sp. 
Inedict. [= unedited, unpublished], Quincy 
9b, 1888 J. Bigot, Mexique. This specimen, 
which is from BMNH, is in poor condition 
and is entirely covered with orange fungus, 
except for the head, which is mounted sep- 
arately on the same pin. 

LECTOTYPE ¢ (here designated to fix 
the current interpretation of this name and 
to ensure stability and uniformity in its fu- 
ture interpretation) of Euphycus setosus: 
(MEI 084117) with the following labels: 
Bilimek, Mexico, 1871, Guadalupe/Cotype 
(pink label)/Phycus setosus Krob., det. 
Kréber 1911/Lectotypus 6, Euphycus se- 
tosus Kroéber, designated in 1998 by SD 
Gaimari et ME Irwin (red label). This 
pinned specimen is in very good condition, 
and is deposited in NHMW. PARALEC- 
TOTYPES: Bilimek, Mexico, 1871/Type 
(pink label)/Phycus setosus Krob., det. 
Kréber 1911/Paralectotypus 2, Euphycus 
setosus Krober, designated in 1998 by SD 
Gaimari et ME Irwin (red label) (pinned °, 
MEI 084118, NHMW); Mexico/Cotype 
(pink label)/Phycus setosus Krob., det. 
Kréber 1911/Paralectotypus 6, Euphycus 
setosus Kroéber, designated in 1998 by SD 
Gaimari et ME Irwin (red label) (2 pinned 
33, MEI 084115, 084116, NHMW); Type 
(pink label)/Mexico, Sta. Fe, 1871/Type 
No. 24183, U.S.N.M. (red label)/Euphycus 
setosus Kroéb., Kréber det. 1911/Paratype 
only, det. WwWirth/Ozodiceromyia mexi- 
cana Bigot, det WWirth/Paralectotypus <6, 
Euphycus setosus Kréber, designated in 


7/8) 


1998 by SD Gaimari et ME Irwin (red la- 
bel) (pinned 3, MEI 032957, USNM). 
Materials examined.—MEXICO: Coa- 
huila, 30.6 km SE of Saltillo, Highway 57, 
2,194.56 m, C. O’Brien, L. O’Brien, G. 
Wibmer, 12-IX-1982 (1 2, MEI 052245, 
MCZC); Distrito Federal, TX-1898 (2 &, 
MEI 032959, 032960, USNM); Mexico 
Citys? R.) Muller iG. <d3MEE032958; 
USNM), W. G. Downs, 14-X-1951 (¢ and 
2, MEI 038775, 038774, AMNH); Pedre- 
gal de San Angel, H. Perez, 27-IX-1969 (1 
2, MEI 084201, KSUC); Tizapan (1 °, 
MEI 052178, AMNH; 2 6, MEI 084194, 
084195, UNAM); Guerrero, 12.9 km E of 
Taxco, 1,828.8 m, R. H. Painter, E. M. 
Painter, 15-[X-1963 (1 6, MEI 052188, 
VAIC); 22 km NE Telolopan, 1,530 m, J. 
A. Powell, J. A. Chemsak, 16-[X-1982 (1 
2, MEI 081567, EMEC); Hidalgo, 16.1 km 
S of Zimapan, 1,889.76 m, J. A. Powell, 
28-IX-1975 (1 3, MEI 052247, SDGC); 
Tepeapulco, G. E. Bohart, W. J. Hanson, 
18-IX-1974 (1 36, MEI 007797, EMUS; 1 
6, MEI 007795, FSCA; 1 3, MEI 007796, 
UAIC; 1 2, MEI 007798, EBCC); Jalisco, 
6.4 km W Mazamilta, 2,072.64 m, R. E 
Smith, 16-X-1950 (1 2, MEI 052182, 
ZMHB); Mexico, C. W. Johnson coll. (1 6, 
MEI 032961, USNM; 1 3, MEI 052180, 
MCZC); Amecameca, R. Dreisbach, K. 
Dreisbach, 25-[X-1957 (1 6, MEI 052192, 
MEIC; 1 2, MEI 052181, INHS); 45.1 km 
SE of San Juan del Rio, G. E. Bohart, W. 
J. Hanson, 31-VIII-1974 (1 6, MEI 
007800, EMUS); Michoacan, 4.8 km W of 
Morelia, 1,859.28 m, R. H. Painter, E. M. 
Painter, 21-X-1963 (1 do, MEI 052185, 
KUIC), 1,950.72 m, R. H. Painter, E. M. 
Painter, 20-[X-1963 (1 6, MEI 052190, 
KUIC; 1 36, MEI 052191, FSCA; 1 2°, MEI 
090233, SDGC); 6 km S of Ocampo, 
Lag.[una] Verde, A. L. Norrbom, 6-X-1991, 
observed waving forelegs acting like a 
wasp (1 2, MEI 052163, USNM); 11.3 km 
E of Quiroga, 2,346.96 m, R. H. Painter, E. 
M. Painter, 21-I[X-1963 (1 ¢, MEI 052176, 
KSUC); 14.5 km W of Ciudad Hidalgo, 
2,194.56 m, R. H. Painter, E. M. Painter, 


580 


19-IX-1963 (1 2, MEI 052177, KSUC); 
14.5 km W of Morelia, 2,072.64 m, J. A. 
Powell, J. A. Chemsak, T. Eichlin, T. P. 
Friedlander, 9-X-1975 (3 3, MEI 052236, 
0522387052239, CASC:'1 6, MEI 052237; 
ZMAS); 20.9 km N of Morelia, 1,981.2 m,, 
R. H. Painter, E. M. Painter, 22-[X-1963 (1 
2, MEI 107043, MEIC); N of Patzcuaro, 3 
km E of Zintzuntzan, 2,100 m, T. Griswold, 
26-X=1987 (1 2, METI. 052229; EMUS); 
SW of Patzcuaro, 3 km E of Zirahuén, 
2,000 m, T. Griswold, 27-X-1987 (1 6, 
MEI 052227, WFBM; 1 2, MEI 052228, 
EMUS); Morelos, 19 km E Cuernavaca, 
Canyon del Lobo, E. M. Fisher, 15-X-1986 
(6 3, MEI 070420, 070421, 070423, 
070424, 070425, 070427, 1 ¢, MEI 
070419, CDFA; 1 6, MEI 070422, 1 2, 
MEI 070428, SDGC; 1 3, MEI 070429, 
ZMUC; 1 6, MEI 070430, MNHN; 1 <4, 
MEI 070431, INHS; 1 2, MEI 070426, 
MEIC; 1 2, MEI 070432, CNCI); Canon 
del Lobos, M. Rodriguez, 19-X-1985 (1 6, 
MEI 084184, CICESE); Tepoztlan, 1,700 
m, C. D. Michener, R. Murillo, J. M. La- 
bougle, 13-XI-1980 (1 36, MEI 052220, 
SEMC); Nuevo Leon, 4.8 km E of Galeana 
Jct., 1,828.8 m, J. A. Chemsak, J. Powell, 
A. Michelbacher, M. M. Michelbacher, 15- 
IX-1976 (1 36, MEI 081532, EMEC; 1 9, 
MEI 081530, NHMW; 1 ¢, MEI 081531, 
SDGC); 6.4 km W Iturbide, 1,676.4 m, J. 
A. Chemsak, 22-IX-1976, at light (1 6, 
MEI 052240, MEIC; 1 2, MEI 052241, 
CASC), J. A. Chemsak, J. A. Powell, 13- 
IX-1976 to 14-IX-1976, at light (1 6, MEI 
052197, EMEC); 8.1 km E of Galeana Jct., 
J. A. Powell, J. A. Chemsak, 16-[X-1976 to 
17-IX-1976 (1 3, MEI 052202, MNHN; 1 
36, MEI 052204, AMNH; 1 6, MEI 
052205.) SEMC; 1 (dé; SME 052206; 
BMNH), J. A. Powell, J. A. Chemsak, A. 
E. Michelbacher, M. M. Michelbacher, 16- 
IX-1976 to 17-IX-1976 (10 o, MEI 
052196, 052199, 052200, 081551, 081550, 
081549, 081548, 081547, 081538, 081537, 
1 ¢&, MEI 081539, EMEC; 1 6, MEI 
052201, UCRC; 1 3, MEI 052198, EBCC; 
1 2, MEI 081536, ZMUC); 11.3 km W of 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Iturbe, 1,767.84 m, R. H. Painter, E. M. 
Painter, 25-[X-1963 (1 6, MEI 052187, 
KSUC); 14.5 km S of junction 60 to Dr. 
Arroyo, R. Turnbow, 22-X-1979 (1 36, MEI 
052174, UGCA); 16.1 km E of San Rob- 
erto, 2,133.6 m, 15-[X-1976 (1 36, MEI 
052203, MZSP); 16.1 km NW Provedencia, 
1828.8 m, J. A. Chemsak, J. Powell, A. 
Michelbacher, M. Michelbacher, 27-IX- 
1976 (6 and 2 in copula, MEI 081553, 
081552, EMEC); 16.1 km N Provedencia, 
J. Powell, J. A. Chemsak, A. Michelbacher, 
M. M. Michelbacher, 25-[X-1976 (1 6, 
MEI 081534, EMEC); 22.5 km N San Juan- 
ito, 2,194.56 m, J. A. Chemsak, J. Powell, 
A. Michelbacher, M. M. Michelbacher, 22- 
IX-1976 (2 3, MEI 081528, 081529, 
EMEC); 66 km S of Saltillo, 1,889.76 m, 
7-IX-1962, U. Kans. Mex. Exped., on flow- 
ers of Encelia farinosa (2 6, MEI 052184, 
052186, KSUC); 66 km SE of Saltillo, 
1,889.76 m, N. Marston, 7-[X-1962 (1 6, 
MEI 052184, KSUC); 70.8 km SE of Sal- 
tillo, G. E. Bohart, W. J. Hanson, 30-VIII- 
1974 (1 3, MEI 007803, EMUS); probably 
Nuevo Leon, Guadalupe, [D.] Bilimek, 
1871 (1 3, 109316, NHMW); Oaxaca, 11.3 
km SE of Nochixtlan, 2,133.6 m, J. A. 
Powell, J. A. Chemsak, T. Eichlin, T. P. 
Friedlander, 7-X-1975 (2 6, MEI 052242, 
052243, CASC; 1 6, MEI 052244, DEIC; 
1 2, MEI 052235, MZSP); 14.5 km NE 
San Jose del Estado, 2,590.8 m, R. E Smith, 
25-X-1966 (1 36, MEI 081533, EMEC); 
San Luis Potosi, 31 km SE San Luis Potosi, 
2,250 m, E. Ramirez, 17-X-1996 (1 3, MEI 
103423, NHMW; 1 6, MEI 103422, 
EBCC; 1 6,-MEI 103420, ZMHB:) ives 
MEI 103419, UCRC), 18-X-1996 (1 ¢, 
MEI 103403, SDGC; 1 3, MEI 103421, 
CNCI); Veracruz, Perote, San Juan del 
Monte, A. Cordoba, 14-X-1989 to 15-X- 
1989 (1 36, MEI 080258, 1 2, MEI 080259, 
IEXA; 1 2, MEI 080260, BMNH); Zaca- 
tecas, 3.2 km S of Luis Moya, 1,920.24 m, 
R. H. Painter, E. M. Painter, 3-[X-1962 (1 
36, MEI 052189, INBC); (unknown state), 
Carr. la para cuautla km 2; 15 Nov. 1978, 
coll: E. Olvara (1 6, MEI 084199, UCDC; 


VOLUME 102, NUMBER 3 


1 3, MEI 084202, UNAM), 19 Nov. 1978 
(1 36, MEI 084200, DEIC); Cuamauaca 
moz, km 57.5 Autopista, 15 Nov. 1978, J. 
C. Medina (1 6, MEI 084188, UCDC); San 
Lucas PUZ., Matorrol espinoso [= spiny 
bush], 21-[X-89, A. Exuihua M. (1 2°, MEI 
079915, UNAM). 

Diagnosis.—The scape in this species is 
densely setose, with long setae. The lower 
frons is globose and distinctly bulging, is 
shiny black, lacking pruinescence, and has 
only a thin line of silver pruinescence along 
the eye margin extending down along the 
parafacial. The face below the antenna is 
shiny black, lacking pruinescence. In males, 
the notum is covered with erect black pile 
interspersed with appressed gold pile. In fe- 
males, the notum and abdominal tergites are 
covered with short, fine, appressed gold 
pile. In the male genitalia, the setae of the 
epandrium and gonocoxites are orange. The 
subepandrial plate is attached to the epan- 
drium only through a thinly membranous 
connection laterally. The sclerotized portion 
of the subepandrial plate is not V-shaped 
posteriorly, but is reduced to two smaller 
lateral plates. The gonocoxites have a su- 
ture along the midline separating the two 
lateral halves. The outer gonocoxal process 
tapers evenly towards apex and is bare of 
setae. The ventral gonocoxal process of the 
gonocoxites is absent; there is a distinct, 
dense clump of orange setae on the poster- 
oventral edge of the gonocoxite. The base 
of the ventral lobes is V-shaped and narrow 
between the two halves. The parameral 
Sheath of the aedeagus has distinct, trans- 
verse wrinkles dorsally. The ventral apo- 
deme lacks a ventral keel. The distiphallus 
is evenly tapered to apex, lacking any 
swelling, and is recurved and parallel with 
the ventral apodeme at the tip. In the female 
terminalia, the furca is notched posteriorly, 
and ventromedially oriented pegs are pres- 
ent on the posterolateral portion of furca. 
The anterolateral furcal prongs are separat- 
ed by a distance subequal to the greatest 
width of the furca. The common sperma- 


581 


thecal duct is not widened basally, and ta- 
pers normally. 

Autapomorphies.—Lower frons globose 
and bulging to antennal insertion (character 
9), posterolateral surface of hindcoxa pilose 
(character 14), parameral sheath with trans- 
verse wrinkless dorsally (character 17), 
subepandrial plate attached to epandrium 
laterally only (character 24), posterolateral 
corners of epandrium held by separate, 
sclerotized membrane, (character 25), pres- 
ence of distinct internal keel where gono- 
coxites fused (character 27), posterior edge 
of furca with distinct notch (character 36), 
medially directed peg present on postero- 
lateral furcal margin (character 37). 

Distribution.—The known distribution 
for this species is found in Fig. 3. Nearly 
all of the specimens were collected at ele- 
vations ranging from 1,500—2,600 m. The 
distribution encompasses parts of the fol- 
lowing biogeographical regions: the Prov- 
incia Xerdfila Mexicana, and the Provincia 
Mesoamericana de Montafia. 

Biology.—This species is most active in 
September and October, with several spec- 
imens collected in late August and in the 
middle of November. Two males were col- 
lected on flowers of Encelia farinosa A. 
Gray (Asteraceae), and one female was ob- 
served in life waving her forelegs in appar- 
ent mimicry of the antennae of a sphecoid 
wasp. 


Ozodiceromyia parargentifera Gaimari 
and Irwin, new species 
(Figs. 36-37, 40) 


Male.—Body length: 7.0 mm. 

Head: 1.04 mm long, 2.00 mm wide, 
1.58 mm high. Distance between eyes at an- 
tennal level 0.62 mm; at genal level 0.83 
mm. Antenna brown, except basal half of 
scape orange. Scape 0.95 mm long, 0.15 
mm wide; with short, fine, black setae, 
evenly distributed over entire scape (includ- 
ing median surface), and few larger setae. 
Pedicel 0.18 mm long, barrel-shaped, setose 
(including median surface). First flagello- 
mere 0.96 mm long, 0.15 mm wide; with 


Fig. 3: 


short setae covering basal two-thirds. Stylus 
0.09 mm long, inserted subapically. Anten- 
nal base to nearest edge of eye 0.15 mm. 
Frons bulging 0.09 mm beyond eye in lat- 
eral view. Face below antenna with silver 
pruinescence. Parafacial lacking pile, with 
silver pruinescence extending to antennal 
base and dorsally along eye margin halfway 
up frons. Frons otherwise shiny black; with 
few, short, fine, black setae (up to 0.17 mm 
long) dorsolateral to antennal base; remain- 
der of frons bare. Genal pile short, dark- 
ened. Palpal pile white basally, dark brown 
distally. Postgenal and occipital pile white; 
occiput with silver pruinescence only along 
edge of eye, and with several black setae. 
Median occipital sclerite flattened; gla- 
brous, shiny black; upper edge not rounded. 
Postocular setae black, arranged in single, 
transverse row. Ocellar tubercle with silver 
pruinescence, and fine, forward-directed, 
black setae. 

Thorax: Scutum and scutellum with fine, 
erect black pile. Scutum 2.22 mm long, 
1.53 mm wide; ground color black; with sil- 
ver-grey pruinescence dorsally, lacking 
pruinescence laterally; median vitta diffuse 
bronze; dc vittae absent; 1 pair dc setae. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Known distribution for Ozodiceromyia mexicana. 


Scutellum with silver pruinescence, reduced 
pruinescence anteriorly. Halter brown. Ka- 
tatergite with dense, long, white pile. Ane- 
pimeron, katepimeron, and meron lacking 
pile, and with reduced silver pruinescence, 
appearing as vertical brown stripe from 
wing base to between second and third cox- 
ae. Anepisternum with silver pruinescence, 
and with white pile. Katepisternum with sil- 
ver pruinescence, with white pile only on 
vertical crest along middle of pleurite. Pre- 
pimeron with silver pruinescence, lacking 
pile. 

Legs: Coxae with silver pruinescence, 
posterior surfaces less so. Posterior surface 
of hindcoxa lacking pile. Forefemur dark 
brown; dorsally with appressed, scale-like 
white pile, and erect, white and brown pile. 
Foretibia with short, erect, black setae; 
proximal, dorsal surface with sparser setal 
covering; orange on basal half, becoming 
dark brown distally; clavate distally. Fore- 
tarsus dark brown. Mid- and hindlegs bro- 
ken off. 

Wing: 5.88 mm long. Basal costal lobe 
with setae arranged in line along outer 
edge, extending into outer row of costal se- 
tae; second line of setae along base of basal 


VOLUME 102, NUMBER 3 


costal lobe, extending into inner row of cos- 
tal setae. Entire membrane slightly dark- 
ened; veins and membrane orange basally, 
darker distally. 

Abdomen: Tergites with erect and re- 
cumbent, white pile; dorsally, with silver 
pruinescence; laterally, lacking pruinesc- 
ence (showing dark brown ground color), 
except tergite 1 and posterior edges of basal 
tergites with silver pruinescence. Sternites 
with silver pruinescence only along anterior 
edge of sternite 2. 

Terminalia: Sternite 8 with fine black se- 
tae restricted to posterior edge, which is 
emarginate medially. Tergite 8 dumbell- 
shaped, with fine black setae restricted to 
posterior edge. Epandrium 0.18 mm long, 
0.68 mm wide at widest point; orange; 
emarginate anteriorly; dorsal surface with 
black setae on posterior half; lateral edges 
parallel; posterolateral corners extended 
and pointed posteriorly. Subepandrial plate 
attached to posterolateral and posterior edg- 
es; sclerotized portion V-shaped posteriorly. 
Sclerotized portion of cerci 0.20 mm long; 
extending posteriorly slightly beyond pos- 
terolateral corners of epandrium; subequal 
in length to ventral epandrial sclerite. Gon- 
ocoxites 0.72 mm wide; orange; with black 
setae, up to 0.47 mm long; fusion 0.36 mm 
long at midline, lacking suture. Inner gon- 
ocoxal process flanged; knob with several 
black setae. Outer gonocoxal process 
flange-like, 0.33 mm long; with short, fine 
setae along dorsal edge. Gonocoxal apode- 
me entirely within anterior edge of gono- 
coxite; lacking sclerotized bridge to para- 
meral sheath of phallus. Ventral gonocoxal 
process 0.27 mm long, flanged; bare. Ven- 
tral lobes distinct, fused basally by thin, 
transparent membrane. Gonostylus expand- 
ed ventrally into large lobe; with dorsobasal 
lobe; with subapical, lateral spur. Dorsal 
apodemes (Fig. 36) of aedeagus parallel; 
shorter than ventral apodeme; parameral 
sheath smooth dorsally. Ventral apodeme 
uniformly wide; with ventral, longitudinal 
keel. Ejaculatory apodeme 0.30 mm long; 
stick-like, expanded distally and bilobed. 


583 


Lateral ejaculatory process a complete ring 
dorsally, but notched; set into aedeagus; 
lightly sclerotized. Distiphallus swollen ba- 
sally, long, undulating, recurved; distally 
perpendicular to ventral apodeme (Fig. 37). 

Female.—Similar to ¢ except as follows: 
body length 9.6 mm. 

Head: 1.22 mm long, 2.28 mm wide, 
1.59 mm high. Distance between eyes at 
level of anterior ocellus 0.50 mm; at anten- 
nal level 0.95 mm; at genal level 1.04 mm. 
Antenna brown. Antennal base to nearest 
edge of eye 0.30 mm. Parafacial pruinesc- 
ence ends at antennal level. Frons shiny 
black; lower frons bulging, with short, fine, 
black setae (up to 0.09 mm long) dorsolat- 
eral to antennal base; upper frons bulging 
and wrinkled, distinct from lower frons. 
Median occipital sclerite and upper edge 
rounded; transverse row of black setae 
across median occipital sclerite in addition 
to row of postocular setae. 

Thorax: Scutum and scutellum with 
erect and recumbent, short, black pile [one 
paratype also has short, fine, appressed gold 
pile]. Scutum 2.58 mm long, 1.76 mm 
wide. Pile of anepisternum and katepister- 
num short, white; densest on upper half of 
anepisternum. 

Legs: Foretibia dark brown [or orange 
basally]. Mid- and hindfemora dark brown. 
Mid- and hindtibiae light brown. Mid- and 
hindtarsi dark brown, except for basal two 
tarsomeres mostly orange. 

Wing: 7.20 mm long. 

Abdomen: Tergites with short, recum- 
bent, white pile. Tergites and sternites with 
reduced silver pruinescence (showing dark 
brown ground color), except posterolateral 
edges of basal tergites with silver pruinesc- 
ence. 

Terminalia (paratype, MEI 027022): 
Furca (Fig. 40) 0.53 mm long, 0.32 mm 
wide; anterior edge not sclerotized; antero- 
lateral prongs dorsoventrally flattened, sep- 
aration narrower than greatest width of fur- 
ca. Furcal bulla not sclerotized; posteriorly 
with indented cavity. Gonopore basal to 
furcal bulla, originating within posterior 


584 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


cavity. Common spermathecal duct basally 
0.11 mm wide, tapering to 0.05 mm within 
0.33 mm distance from gonopore; 0.62 mm 
long. Spermathecal ducts originate from 
clean trifurcation with central sac duct; cen- 
tral sac duct wider in basal diameter than 
spermathecal ducts. Central sac duct 0.45 
mm long. Central sac 0.68 mm long, 0.41 
mm wide. Spermatheca 0.17 mm in diam- 
eter; rounded, but basal edge slightly flat- 
tened. 

Type materials—HOLOTYPE 36 (MEI 
027017) with the following label: MEXI- 
CO, Guerrero, 4 mi. W of Chilpancingo, 
July 15, 1984, Carroll, Schaffner, Friedlan- 
der. This pinned specimen, which is depos- 
ited in CASC (with permission of TAMU), 
is in good condition except for its missing 
mid- and hindlegs. 

Materials examined.—PARATYPES. 
MEXICO: same label as holotype (1 &, 
MEI 027022, SDGC); Michoacan, 9.3 km 
SE Quiroga, 25-VIII-1991, W. FE Barr, on 
roadside vegetation, 2,653 m (1 2, MEI 
079906, WFBM); Puebla, 4.4 mi. SW [San 
Francisco] Acatepec, 9-VII-1981, Bogar, 
Schaffner, Friedlander (allotype 2, MEI 
027003, TAMU). 

Diagnosis.—This species is much like 
Ozodiceromyia argentifera and Ozodicero- 
myia livdahli. The scape in this species is 
setose, but most of the setae are short, with 
few longer ones. The lower frons bulges 
only slightly, with silver pruinescence lat- 
erally, extending down along parafacial. 
The face below the antenna also has silver 
pruinescence. In males, the notum has some 
fine, erect, black setae (the holotype, the 
only known male, appears rubbed dorsally, 
and so presence or condition of appressed 
pile is unknown). In females, the notum has 
short, fine, erect and appressed black pile, 
and occasionally appressed gold pile. In the 
male genitalia, the setae of the epandrium 
and gonocoxites are black. The subepandri- 
al plate is attached to the epandrium at the 
posterolateral corners, and the sclerotized 
portion of the subepandrial plate is V- 
shaped posteriorly. The gonocoxites lack a 


suture along the midline separating the two 
lateral halves. The outer gonocoxal process 
is widened distally, with a small patch of 
setae on the dorsolateral surface. The ven- 
tral gonocoxal process of the gonocoxites is 
present, elongated and flattened; there are 
no clumped setae at its base. The base of 
the ventral lobes is U-shaped and broad be- 
tween the two halves. The parameral sheath 
of the aedeagus is smooth dorsally. The 
ventral apodeme has a ventral keel. The dis- 
tiphallus is swollen basally, and is perpen- 
dicular to the ventral apodeme throughout. 
In the female terminalia, the furca is not 
notched posteriorly, and lacks a posterolat- 
eral peg. The anterolateral furcal prongs are 
separated by a distance narrower than the 
greatest width of the furca. The common 
spermathecal duct is broad basally, at half 
the width of the entire furca; the duct tapers 
quickly. 

Autapomorphies.—Halter brown (char- 
acter 15), ventral apodeme longer than dor- 
sal apodemes (character 18), ventral surface 
of ventral apodeme with longitudinal keel 
(character 19), distiphallus perpendicular to 
dorsal apodemes (character 23, state 0), an- 
terior prongs of furca narrowly separated 
(character 38). 

Distribution.—The known distribution 
for this species is found in Fig. 4. This spe- 
cies is found in southern Mexico, in the 
states of Puebla, Michoacan, and Guerrero. 
The type locality, in Guerrero, is part of the 
northern slope foothills of the Sierra Madre 
de Sur range, at an elevation over 1,500 m. 
The remaining localities are all above this 
elevation, as high as 2,600 m. The distri- 
bution encompasses parts of the following 
biogeographical regions: the Provincia Me- 
soamericana de Montana, and the Provincia 
Pacifica. 

Etymology.—Gr., para: beside, near + 
“argentifera,’ referring to the similarity 
with Ozodiceromyia argentifera (Krober); 
to be treated as a noun in apposition. 

Biology.—Adults are active at least 
through the months of July and August. 


VOLUME 102, NUMBER 3 


585 


Fig. 4. 


KEY TO SPECIES 


A comprehensive key to species of Ozo- 
diceromyia is not presented herein. This 
will be included in the larger revision of the 
entire genus in progress by the authors. The 
current key is a continuation of Gaimari and 
Irwin’s (2000) key to world genera of Cy- 
clotelini. An alternative is the key to the 
therevid genera of North America found in 
Irwin and Lyneborg (198la, b). The first 
couplet below separates members of the 


zodiceromyia signatipennis 


Ozodiceromyia costalis 


2 4 10 11 21 29 31 32 35 


1 5 6 7 8 15 16 20 23 33 34 39 


Ripe: 


3 11 22 26 28 29 30 35 40 41 


Known distribution for Ozodiceromyia parargentifera. 


Ozodiceromyia mexicana-group from. all 
other members of the genus. Note that the 
male genitalic characters in the key are usu- 
ally visible without dissection. The female 
genitalic character in couplet 3 requires dis- 
section, but should not be necessary for 
routine identification. 


il. Antenna (Figs. 7-10) longer than head. 
Scape setose throughout. Pedicel barrel- 
shaped, longer than wide, setose throughout. 
First flagellomere setose over basal two- 


9 14 17 24 25 27 36 37 


zodiceromyia mexicana 
15 18 19 23 38 
zodiceromyia parargentifera 


zodiceromyia argentifera 


Ozodiceromyia livdahli 


Single most parsimonious cladogram for the species of the Ozodiceromyia mexicana-group, showing 


character state changes under ACCTRAN character optimization. Characters are numbered as in the text; hash 
marks are as follows: black = forward change with no homoplasy; white = change with homoplasy. 


586 


O. signatipennis 


O. costalis 


O. mexicana 


O. parargentifera 


O. argentifera 


O. livdahli 


Single most parsimonious cladogram for 


Fig. 6. 
the species of the Ozodiceromyia mexicana-group, 
with Bremer support indices indicated above each 
branch and post-successively reweighted, rescaled Bre- 
mer support indices below each branch. 


thirds. Wing membrane and veins orange, at 
least basally; distal third of wing often dark- 
GneGenwithudankeneGuvielisS mei elie eect 
SiS Saris |e eee Ozodiceromyia mexicana-group 
= Antenna (Fig. 11) shorter than head, rarely 
as long as head. Scape usually bare on me- 
dian surface. Pedicel as long as wide, usu- 
ally bare on median surface. First flagello- 
mere setose at most in basal third. Wing 
membranes hyaline or banded, but never or- 
ange; veins dark, rarely orange; distal third 
of wing not darkened relative to basal two- 
thirds other Ozodiceromyia 
2(1). Scape densely setose, with long setae, lon- 
ger than those of first flagellomere (Figs. 9— 
10). Lower frons globose, distinctly bulging, 
and shiny black, with silver pruinescence 
only along eye margin (Figs. 9-10). Face 
beneath antenna shiny black, lacking prui- 
nescence (Figs. 9-10). Male: notum with 
erect black pile; posterior surface of hind- 
coxa with long, white pile (Fig. 16); outer 
gonocoxal process tapering evenly, bare 
(Fig. 31); ventral gonocoxal process of gen- 
italia absent, dense clump of orange setae 
on corresponding edge (Fig. 30) 
Ozodiceromyia mexicana 
— Scape setose, but with short setae, with few 
long setae interspersed (Figs. 7-8). Lower 
frons not globose, only slightly bulging; sil- 
ver pruinescence present dorsolaterally to 
antennal base (Figs. 7-8). Face beneath an- 
tenna with silver pruinescence (Figs. 7-8). 
Male: pile of notum variable, but usually 
both erect and appressed pile present; pos- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


terior surface of hindcoxa lacking pile (Fig. 
14); outer gonocoxal process widened dis- 
tally, with small patch of short setae on dor- 
solateral surface (Fig. 27); ventral gonocox- 
al process present, long and flattened, with 
no densely clumped setae (Fig. 26) 
. Halter brown in both sexes. Male: notum 
with erect black pile; setae of gonocoxites 
and epandrium black; distiphallus perpen- 
dicular to ventral apodeme throughout 
length (Fig. 37). Female: distance between 
anterolateral furcal prongs distinctly nar- 
rowed anteriorly (Fig. 40) 
waite ay fe ae WA gee Ozodiceromyia parargentifera 
- Halter yellow in both sexes. Male: notum 
with both erect and appressed pale or gold 
pile; setae of gonocoxites and epandrium 
black or orange; distiphallus recurved be- 
fore apex to become parallel with ventral 
apodeme (Fig. 33). Female: distance be- 
tween anterolateral furcal prongs not nar- 
rowed anteriorly, subequal to widest part of 
furca (Fig. 39) 
4(3). Male: setae of gonocoxites and epandrium 
black (Figs. 23, 26—27). Female: appressed 
pile on abdominal tergites gold 


Ozodiceromyia argentifera 
— Male: setae of gonocoxites and epandrium 
pale or orange (Fig. 28). Female: appressed 
pile on abdominal tergites brown and white 
SACRE es ah ace Recker Ozodiceromyia livdahli 


CHARACTER DESCRIPTIONS 


This section is devoted to description of 
morphology and taxon distribution of states 
of the 41 characters used in this analysis. 
Of these, 11 are taken from the head, 5 
from the thorax, 19 from the male termin- 
alia, and 6 from the female terminalia. 
Characters described and discussed by Gai- 
mari and Irwin (2000) are referred to the 
appropriate character number therein for 
description, although state distributions are 
discussed herein. All characters are binary. 
Character distributions are discussed rela- 
tive to the presented cladogram (Fig. 5). 


HEAD 
I. Antennal length 


0 shorter than or subequal to head 
length 
1 longer than head 
The states for this character are described 
as character 1 in Gaimari and Irwin (2000). 


VOLUME 102, NUMBER 3 587 


Figs. 7-13. Heads, ¢ (7, 9, 11), 2 (8, 10); 2, antennal first flagellomere (12), tip only (13). (7-8) Ozodi- 
ceromyia argentifera (MEI 038790, 052183). (9-10, 13) Ozodiceromyia mexicana (MEI 052192, 107043, 
070426). (11-12) Ozodiceromyia signatipennis (MEI 044750, 056973). Measure bars, 0.2 mm. 


In the current analysis, an elongated anten- 2. Medial surface of scape 
na (Fig. 7) is autapomorphic for the mexi- 

cana-group, and is found in no other known 0 without setae 
Ozodiceromyia. Members of several other 1 setose 


cycloteline genera, including Cyclotelus In nearly all Ozodiceromyia, the scape is 
Walker, also share this state. bare on the median surface (Fig. 11). Only 


588 


in the mexicana-group and the outgroup 
taxon Ozodiceromyia costalis (and several 
undescribed, putatively related species) is 
this surface setose (Fig. 9). 


3. Setae of scape 


0 longer than those of first flagellomere 
1 most are shorter than or equal to 
those of first flagellomere 

The setae of the scape are longer than those 
of the first flagellomere (Fig. 9), plesiom- 
orphically, as in both outgroup taxa and 
Ozodiceromyia mexicana. As a Synapomor- 
phy for the clade of Ozodiceromyia parar- 
gentifera + Ozodiceromyia argentifera + 
Ozodiceromyia livdahli, the setae of the 
scape are shorter than or equal to those of 
the first flagellomere, at most with several 
longer setae interspersed among the short 
setae (Fig. 7). 


4. Medial surface of pedicel 


Q without setae 

lL Ssetose 
The states for this character are described 
as character 3 in Gaimari and Irwin (2000). 
The apomorphically setose medial surface 
of the pedicel (Fig. 9) is present in all mem- 
bers of the mexicana-group, and in the out- 
group taxon Ozodiceromyia costalis. 


5. Setae of first flagellomere 


QO basal only, or bare 

1 covering more than basal half 
In nearly all Ozodiceromyia, including both 
outgroup taxa, the setae of the first flagel- 
lomere are restricted to the base (Fig. 11). 
Only as an autapomorphy for the mexicana- 
group do these setae cover more than the 
basal half of the first flagellomere (Fig. 9). 


6. Setae of first flagellomere 


OQ less dense ventrally, or bare 

1 fully surrounding with equal density 
Plesiomorphically, the ventral surace of the 
first flagellomere is bare, or only sparsely 
setose (Fig. 11), as in nearly all Ozodicer- 
omyia including both outgroup taxa. Only 
as an autapomorphy for the mexicana- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


group do setae fully surround the first fla- 
gellomere with equal density (Fig. 9). 


7. Antennal style 


QO originates near tip, but extended 
beyond first flagellomere distally 
1 distinctly subapical, and not extended 
beyond tip of first flagellomere 
Plesiomorphically, the antennal style origi- 
nates very close to the apex of the first fla- 
gellomere, but slightly subapical and ven- 
tral. Despite this, the tip of the style extends 
distally beyond the tip of the first flagellom- 
ere (Fig. 12). As an autapomorphy for the 
mexicana-group, the antennal style origi- 
nates distinctly subapically on the first fla- 
gellomere within a ventral pit, with the sty- 
lar tip not reaching the apex of the first fla- 
gellomere (Fig. 13). 


8. First stylar segment 


0 as long as wide 

1 wider than long, and ring-like 
Plesiomorphically, the first stylar segment 
is as long as wide (Fig. 12). As an auta- 
pomorphy for the mexicana-group, this seg- 
ment is reduced to a small, flattened ring 
set within a ventral pit (Fig. 13). 


9. Lower frons at antennal insertion 


O normal, not globose 

1 globose, bulging 
At the antennal insertion, the lower frons 
bulges slightly in most Ozodiceromyia (Fig. 
7). As an autapomorphy for Ozodiceromyia 
mexicana, the lower frons is distinctly bulg- 
ing and globose (Figs. 9-10). 


10. Setae of 3 frons 


O scattered 

1 in patches or absent 
In many Ozodiceromyia, including the out- 
group taxon Ozodiceromyia signatipennis, 
the setae on the male frons are scattered 
(Fig. 11). In the mexicana-group and Ozo- 
diceromyia costalis, these setae are absent 
or reduced to small patches located dorso- 
laterally from the antennal base (Fig. 7). 


VOLUME 102, NUMBER 3 


589 


Figs. 14-16. 
of wing, @, left is basal (15); hindcoxa, d, left is anterior (16). (14-15) Ozodiceromyia argentifera (MEI 
037789, 052183); arrow (14) points to katepisternum. (16) Ozodiceromyia mexicana (052192). Measure 
bars, 0.2 mm. 


11. Face directly below antenna 


QO with full pruinescence 
1 shiny, with little or no pruinescence 
(usually black) 
For most Ozodiceromyia, the face below 
the antenna is fully covered with silver 


Thorax and associated structures: lateral thorax, d, left is anterior (14); basal costal lobe 


pruinescence (Fig. 8). In Ozodiceromyia 
costalis (and several undescribed, putative- 
ly related species) and Ozodiceromyia mex- 
icana, the face completely lacks pruinesc- 
ence, or any other vestiture, exposing the 
black, shiny cuticle (Fig. 9). 


590 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 17-25. 4d genitalic and pregenitalic structures: 8th sternite (17) and tergite (18); gonostylus (19-20); 
ventral lobe (21—22); epandrium, split view (left, dorsal; right, ventral) (23, 25), dorsal view (24). (17-19, 23) 
Ozodiceromyia argentifera (MEI 037789); arrow (23) points to subepandrial plate. (20, 22) Ozodiceromyia 
signatipennis (MEI 044750). (21, 25) Ozodiceromyia mexicana (MEI 052240); arrow (21) points to pilose, 
secondary distal lobe. (24) Ozodiceromyia livdahli (MEI 038822). Measure bars, 0.2 mm. 


VOLUME 102, NUMBER 3 


THORAX 
12. S notal vestiture 


OQ without decumbent pile 

1 with thick decumbent pile 
Although decumbent pile on the male no- 
tum is present in other members of this ge- 
nus, and other therevids, its absence is con- 
sidered plesiomorphic at this level of anal- 
ysis. The presence of thick, decumbent pile 
is a synapomorphy for Ozodiceromyia ar- 
gentifera + Ozodiceromyia livdahli. 


13. Fine, erect setae of 3 notum 


O some black 

1 entirely pale or gold, none black 
All taxa studied have fine, erect setae on 
the male notum. Plesiomorphically, most of 
these setae are black, although some pale 
setae may be interspersed. As a synapo- 
morphy for Ozodiceromyia argentifera + 
Ozodiceromyia livdahli, all of these fine, 
erect, notal setae are pale or gold. Note, this 
does not include the notal macrosetae. 


14. Posterolateral surface of hindcoxa 


0 lacking pile 

1 pilose 
Plesiomorphically, the posterolateral sur- 
face of the hindcoxa is bare (Fig. 14). As 
an autapomorphy for Ozodiceromyia mexi- 
cana, this surface has long, thin pile (Fig. 
16). This state is also present in certain oth- 
er groups within Ozodiceromyia. 


15. Halter color 


QO brown 

1 yellow 
In most species of Ozodiceromyia, the hal- 
ter is brown. In several groups, including 
the mexicana-group, yellow halter is syna- 
pomorphic. The only exception is the 
brown halter found in Ozodiceromyia par- 
argentifera, which is considered an auta- 
pomorphy for the species. 


16. Wing color 


O clear, or with darkened patches 
1 cloudy yellow or orange 


591 


Although banded wings are known in Ozo- 
diceromyia, clear wings or those with dark- 
ened patches along certain wing veins are 
plesiomorphic at this level of analysis. Only 
in the mexicana-group is the wing entirely 
cloudy yellow or orange, becoming dark- 
ened in the distal third. This is considered 
an autapomorphy for the mexicana-group. 


MALE TERMINALIA 
17. Parameral sheath texture 


OQ smooth dorsally 

1 with transverse wrinkles dorsally 
Plesiomorphically, the parameral sheath of 
the aedeagus is smooth dorsally (Fig. 32). 
As an autapomorphy for Ozodiceromyia 
mexicana, the parameral sheath has distinct 
transverse wrinkles dorsally (Fig. 34). 


18. Relative lengths of dorsal and ventral 
apodemes 


QO subequal, or dorsal apodeme longer 
1 ventral apodeme longer 
The states for this character are described 
as character 37 in Gaimari and Irwin 
(2000). The longer ventral apodeme (Fig. 
37) is autapomorphic for Ozodiceromyia 
parargentifera. 


19. Ventral surface of ventral apodeme 


O without longitudinal keel 

1 with longitudinal keel 
Most species of Ozodiceromyia lack a ven- 
tral longitudinal keel on the ventral apode- 
me. A longitudinal keel is present (Fig. 37) 
as an autapomorphy for Ozodiceromyia 
parargentifera. 


20. Distiphallus length 


0 shorter than dorsal apodemes 
1 as long or longer than dorsal apo- 
demes 

At this level of analysis, having a disti- 
phallus shorter than the dorsal apodemes is 
considered plesiomorphic (Fig. 38). As an 
autapomorphy for the mexicana-group, the 
distiphallus is distinctly longer than the dor- 
sal apodemes (Fig. 33). 


592 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


VOLUME 102, NUMBER 3 


21. Dorsal part of basal distiphallus 


QO with median groove 

1 without median groove, rounded 
A median, longitudinal groove is present on 
the dorsal part of the basal distiphallus in 
the outgroup taxon Ozodiceromyia signati- 
pennis. This groove is lacking, and the dis- 
tiphallus is entirely rounded above in the 
mexicana-group and Ozodiceromyia costal- 
is. 

22. Basal portion of distiphallus 


Q tapering evenly 

1 swollen 
Plesiomorphically, the basal portion of the 
distiphallus is not swollen and tapers grad- 
ually and evenly to the apex (Fig. 35). As 
a synapomorphy for Ozodiceromyia parar- 
gentifera + Ozodiceromyia argentifera + 
Ozodiceromyia livdahli, this part of the dis- 
tiphallus is distinctly swollen (Figs. 33, 37). 


23. Orientation of distiphallus at apex 


Q perpendicular to dorsal apodemes 

1 parallel with dorsal apodemes 
In both outgroup taxa, the distiphallus is 
perpendicular to the dorsal apodemes 
throughout, not recurving anteriorly or 
posteriorly. As a synapomorphy for mem- 
bers of the mexicana-group, the distiphal- 
lus recurves anteriorly before its apex, be- 
coming parallel with the dorsal apodemes 
(Figs. 33, 35). As an autapomorphy for 
Ozodiceromyia parargentifera, the disti- 
phallus loses its curve and becomes per- 
pendicular with the dorsal apodemes (Fig. 
37); as in state 0: 


24. Subepandrial plate attachment to 
epandrium 


Q laterally and posteriorly 
1 laterally only 


ee 


Figs. 26-31. 


595 


Plesiomorphically, the subepandrial plate is 
solidly attached to the epandrium both lat- 
erally and posterolaterally (Fig. 23). As an 
autapomorphy for Ozodiceromyia mexi- 
cana, the subepandrial plate is greatly re- 
duced, and is only attached to the epan- 
drium through a lateral membranous con- 
nection (Fig. 25). 


25. Posterolateral corners of epandrium 


O held by subepandrial plate 

1 held by sclerotized membrane 
The posterolateral corners of the epandrium 
are held together by the subepandrial plate 
(Fig. 23), plesiomorphically. As an auta- 
pomorphy for Ozodiceromyia mexicana, 
the corners are held by a separate, sclero- 
tized membrane (Fig. 25), apparently not 
associated with the subepandrial plate. 


26. Gonocoxal setae color 


O mostly pale 

1 all black 
Although this is variable throughout Ozo- 
diceromyia, pale gonocoxal setae are ple- 
siomorphic at this level of analysis. Under 
the accelerated transformation character op- 
timization model, presence of all black se- 
tae (Fig. 27) is synapomorphic for the clade 
of Ozodiceromyia parargentifera + Ozo- 
diceromyia argentifera + Ozodiceromyia 
livdahli, with a subsequent autapomorphic 
change to pale setae in Ozodiceromyia li- 
vdahli (Fig. 28) 


27. Distinct internal keel where 
gonocoxites fused 


QO absent 

1 present 
At this level of analysis, the lack of a dis- 
tinct internal keel (visible externally as a 
groove) where the gonocoxites are fused 


3d gonocoxites and associated structures, split view (left, ventral; right, dorsal) (26, 30), lateral 


view, gonostylus removed, left is dorsal (27), lateral view, left is dorsal (28-29, 31). (26-27) Ozodiceromyia 
argentifera (MEI 037789); arrow (26) points to ventral gonocoxal process, arrow (27) points to inner gonocoxal 
process. (28) Ozodiceromyia livdahli (MEI 038822); arrow points to outer gonocoxal process. (29) Ozodicero- 
myia signatipennis (MEI 044750). (30-31) Ozodiceromyia mexicana (MEI 052240). Measure bars, 0.2 mm. 


594 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 32-38. Aedeagal complex: dorsal view (32, 34, 36); lateral view (33, 35, 37-38). (32-33) Ozodicer- 
omyia argentifera (MEI 037789); arrow (32) points to lateral ejaculatory process, arrow (33) points to disti- 
phallus. (34-35) Ozodiceromyia mexicana (MEI 052240); arrow (34) points to ejaculatory apodeme, arrow (35) 
points to dorsal apodeme. (36-37) Ozodiceromyia parargentifera (MEI 027017); arrow (37) points to ventral 
apodeme. (38) Ozodiceromyia signatipennis (MEI 044750). Measure bar, 0.2 mm. 


VOLUME 102, NUMBER 3 


(Fig. 26) is plesiomorphic. As an autapo- 
morphy for Ozodiceromyia mexicana, this 
distinct line of fusion is present (Fig. 30). 


28. Ventral gonocoxal process 


Q absent or only small fold of edge 

1 elongated and flanged 
Plesiomorphically, the ventral gonocoxal 
process is absent, or is only a small fold of 
the edge (Fig. 30). As a synapomorphy for 
the clade of Ozodiceromyia parargentifera 
+ Ozodiceromyia argentifera + Ozodicer- 
omyia livdahli, the ventral gonocoxal pro- 
cess is present, elongated, and flanged (Fig. 
26). 


29. Setae of ventral gonocoxal process 
(including corresponding edge if absent) 


QO absent 
1 distinctly different 
gonocoxal setae 

Plesiomorphically, setae are absent on the 
ventral gonocoxal process (Fig. 26), or on 
the corresponding edge if absent. In Ozo- 
diceromyia mexicana and Ozodiceromyia 
costalis, the setae are distinctly different 
from the remaining gonocoxal setae (Fig. 
30). 


from other 


30. Setae of outer gonocoxal process 


Q absent 

1 present 
The outer gonocoxal process is bare of se- 
tae (Fig. 31) plesiomorphically. As a syna- 
pomorphy for the clade of Ozodiceromyia 
parargentifera + Ozodiceromyia argenti- 
fera + Ozodiceromyia livdahli, there is a 
small patch of short setae on the dorsolat- 
eral portion of the outer gonocoxal process 
(Fig. 27). 


31. Outer and ventral gonocoxal 
processes 


O fused 

1 unfused 
The outer and ventral gonocoxal processes 
are plesiomorphically fused (Fig. 29), as in 
the outgroup taxon Ozodiceromyia signati- 
pennis. These processes are separate and 


595 


unfused (Figs. 27, 35) in the mexicana- 
group and in Ozodiceromyia costalis. 


32. Projection on gonocoxal apodeme for 
articulation with aedeagus 


QO absent 

1 present 
There is no sclerotized connection between 
the gonocoxal apodeme and the aedeagus 
in any taxon in this study. Ozodiceromyia 


signatipennis completely lacks any projec- 


tions on the gonocoxal apodeme. However, 
in the mexicana-group and in Ozodicero- 
myia costalis, there is a medially directed 
projection of the gonocoxal apodeme (Fig. 
30) that provides a site for articulation with 
the aedeagus. 


33. Distal part of ventral lobe 


OQ bare, or with fine, short pile on outer 
edge 

1 with pilose distal lobe 
Most Ozodiceromyia, including both out- 
group taxa, lack a secondary distal lobe on 
the ventral lobe (Fig. 22). Only as an au- 
tapomorphy for the mexicana-group is a 
finely pilose, secondary lobe present distal- 
ly on the ventral lobe (Fig. 21). 


34. Lobe of gonostylus 


Q expanded ventrally into small lobe 

1 lobe greatly expanded ventrally 
In the more primitive Ozodiceromyia (see 
Gaimari 1998), the gonostylus is strap-like, 
with no lobe ventrally. More advanced 
members of the genus have a small lobe on 
the ventral part of the gonostylus (Fig. 20), 
which is the plesiomorphic state at this lev- 
el of analysis. As an autapomorphy for the 
mexicana-group, this lobe is greatly en- 
larged (Fig. 19). 


35. Subapical spur of gonostylus 


OQ present 

1 absent 
The states of this character are described 
as character 56 in Gaimari and Irwin 
(2000). The subapical spur of the gono- 
stylus is absent in both Ozodiceromyia 


596 


mexicana and the outgroup taxon Ozodi- 


ceromyia costalis. Although this character 


is quite variable in the genus, the change 
to state 0 appears to be a synapomorphy 
for the clade of Ozodiceromyia parargen- 
tifera + Ozodiceromyia argentifera + 


Ozodiceromyia livdahli. 
FEMALE TERMINALIA 


36. Posterior edge of furca 


Q rounded 

1 with notch or bend 
Plesiomorphically, the posterior edge of the 
furca is rounded normally, with no notches 
or bends (Fig. 39). As an autapomorphy for 
Ozodiceromyia mexicana, a distinct, ante- 
riorly directed notch is present (Fig. 41). 


37. Medioventral edge of posterolateral 
furca 


QO smooth 

1 with small process 
Plesiomorphically, there are no medially di- 
rected pegs on the inner furcal margin (Fig. 
39). As an autapomorphy for Ozodicero- 
myia mexicana, there is a small process, or 
peg, on the inner ventral edge of the furca 
on the posterolateral margin (Fig. 41). 


38. Space between anterior prongs of 
furca 


0 subequal to greatest furcal width 
1 distinctly narrower than greatest 
furcal width 

The space between the anterior prongs of 
the furca is approximately the same width 
as the greatest furcal width (Fig. 39), ple- 
siomorphically. As an autapomorphy for 
Ozodiceromyia parargentifera, the width 
between the anterior prongs of the furca is 
distinctly narrowed (Fig. 40). 


39. Posterior part of furcal bulla 


QO normal, not indented 
1 indented as cavity to house common 
spermathecal duct 
Plesiomorphically, the posterior part of the 
furcal bulla us rounded normally. As an au- 
tapomorphy for the mexicana-group, this 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


posterior margin is distinctly indented as a 
cavity to house the common spermathecal 
duct (Fig. 39). 


40. Basal part of common spermathecal 
duct 


QO normal width, more or less even 
throughout length 

1 distinctly widened, tapering quickly 
The basal part of the common spermathecal 
duct is plesiomorphically of normal width, 
more or less even throughout its length, 
only gradually tapering (Fig. 41). As a syn- 
apomorphy for the clade of Ozodiceromyia 
parargentifera + Ozodiceromyia argenti- 
fera + Ozodiceromyia livdahli, the base is 
distinctly enlarged and widened, tapering 
quickly to normal width (Fig. 39). 


41. Individual spermathecal duct origin 


Q from base of central sac 

1 from common spermathecal duct 
Plesiomorphically, the spermathecal ducts 
originates from the base of the central sac 
(Fig. 41). As a synapomorphy for the clade 
of Ozodiceromyia parargentifera + Ozo- 
diceromyia argentifera + Ozodiceromyia 
livdahli, the spermathecal ducts originate 
on the common spermathecal duct, with a 
short duct then leading to the central sac 
(Fig. 39). 


RESULTS AND DISCUSSION 


Cladistic analysis.—The analysis pro- 
duced a single most parsimonious tree (Fig. 
5) of length 47, Consistency Index (Kluge 
and Farris 1969) of 0.87 (0.80 excluding 
autapomorphies), Retention Index (Farris 
1989) of 0.81, and Rescaled Consistency 
Index (Farris 1989) of 0.71. The topology 
remained stable through successive weight- 
ing (Farris 1969; Carpenter 1988, 1994), 
implemented by PAUP by reweighting all 
characters on a base weight of 1,000 by the 
maximum values of their Rescaled Consis- 
tency Indices, leveling out at a single iter- 
ation. After this single iteration, the tree sta- 
tistics above were all 0.97 or higher. Be- 
cause the expected amount of homoplasy 


VOLUME 102, NUMBER 3 


S27) 


Figs. 39-41. @ reproductive structures: furca (dorsal view) through central sac and basal part of sper- 
mathecal ducts (both spermathecal ducts truncated basally) (39); furca (dorsal view) with truncated common 
spermathecal duct (40); furca (dorsal view) through central sac and spermathecae (41). (39) Ozodiceromyia 
argentifera (MEI 052230); arrow points to central sac. (40) Ozodiceromyia parargentifera (MEI 027022); 
arrow points to furcal bulla. (41) Ozodiceromyia mexicana (MEI 070426); arrow points to spermatheca. 


Measure bars, 0.2 mm. 


increases with the number of taxa in a par- 
simony analysis, the regression equation of 
Sanderson and Donoghue (1989) was used 
to calculate the expected value for the Con- 


sistency Index. The calculated value of 0.78 
for 6 taxa is lower than the value indicated 
in the current analysis, excluding autapo- 
morphies, so the amount of homoplasy is 


598 


less than expected for an analysis with this 
number of taxa under the 1989 model. The 
relatively high Retention Index indicates 
that a high proportion of the potential syn- 
apomorphies in the data matrix are present 
as homologies on the cladogram. 

Bremer support (aka, decay analysis) 
may reflect the robustness of certain ele- 
ments of the resulting cladograms (Bremer 
1988, Donoghue et al. 1992, Kallersj6 et al. 
1992). A decay analysis entails calculating 
the difference in tree length between the 
most parsimonious tree and the shortest tree 
which lacks each nodal group. This reflects 
how much additional evidence, in the form 
of characters supporting an alternative 
grouping, would be necessary to overthrow 
a clade in the most parsimonious hypothe- 
sis. Using AUTODECAY (Eriksson 1997, 
currently in version 2.9.9) in combination 
with PAUP, Bremer support indices were 
calculated for each nodal group in the clad- 
ogram. Following Gaimari and Irwin 
(2000), Bremer support indices were also 
calculated using the successively reweight- 
ed data, and were rescaled per the proce- 
dure outlined by Bremer (1994). The re- 
sulting Bremer support indices for each 
node are displayed on the cladogram in Fig. 
6. The Bremer support values are an indi- 
cation of the number of characters in op- 
position to the current topology that would 
be necessary to break down the individual 
nodes of the tree. Justification for excluding 
perturbation (e.g., bootstrap, jackknife) and 
permutation (PTP, T-PTP) based methods to 
assess branch support is provided by Gai- 
mari and Irwin (2000). 

Classification.—The internal classifica- 
tion of the Therevidae has only recently be- 
gun to receive attention. Not until recently 
(Irwin and Lyneborg 1981a) was the family 
formally divided into two subfamilies, 
Therevinae and Phycinae. More recently, 
Gaimari and Irwin (2000) defined and pro- 
posed a phylogenetic classification for gen- 
era of Cyclotelini, including Ozodicero- 
myia, and Gaimari (1998) defined putative 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 3. Classification for the species of the Ozo- 
diceromyia mexicana-group. 


Cyclotelini Gaimari and Irwin, 2000 
Ozodiceromyia Bigot, 1890 
Ozodiceromyia **mexicana-group’’ 
O. mexicana Bigot, 1890 
O. parargentifera, n. sp. 
O. argentifera (Krober 1929) 
O. livdahli, n. sp. 


species-groups within the genus, including 
the mexicana-group as defined herein. 

According to the cladogram, the Bremer 
support index of 7 additional steps for the 
clade, and the rescaled Bremer support in- 
dex of 13 additional steps, the Ozodicero- 
myia mexicana-group appears well support- 
ed by autapomorphies. This hypothesis will 
be most thoroughly tested in the phyloge- 
netic analysis of the entire genus underway. 
The classification in Table 3 is presented as 
a phyletic sequence of taxa (Nelson 1969, 
1972, 1973) within this small group of Ozo- 
diceromyia. The group will be classified 
within the larger context of the entire genus 
with the pending revisionary work under- 
way. 


ACKNOWLEDGMENTS 


Heartfelt thanks are extended to Jill D. 
Mullett for her superb illustrations; Gail E. 
Kampmeier (INHS) (and our cadre of lab 
assistants, including Mona Abou-Batnin, 
Jessica Adams, Jamie Bender, Jessica Bick- 
ham, Amanda Buck, Shelly Cook, Andrea 
Gavurnik, Mary Grabowski, Dyanna Greg- 
ory, Traci Gustafson, Sarah Kastelic, Sandy 
Kawanaka, Erin Leslie, Leslie Marsh, Rob- 
ert Pollok, Ann Ramsey, Stephanie Stroth- 
off, Johnson Zeledon, and Anjun Zhou) for 
making this research much easier through 
databasing; Martin Hauser (INHS), Kevin 
C. Holston (INHS), the late Ellis G. 
MacLeod (University of Illinois), Mark A. 
Metz (INHS), Evert I. Schlinger (Schlinger 
Foundation), Donald W. Webb (INHS), Bri- 
an M. Wiegmann (North Carolina State 
University), Shaun L. Winterton (UQIC), 


VOLUME 102, NUMBER 3 


Long-Long Yang (North Carolina State 
University), and David K. Yeates (Univer- 
sity of Queensland), for many fruitful dis- 
cussions about systematic theory, therevid 
and asiloid evolution and characters, and 
for numerous shared experiences in the 
field. We also thank Neal L. Evenhuis 
(BPBM) and E Christian Thompson (Sys- 
tematic Entomology Lab, USDA) for pro- 
viding helpful nomenclatural and _ biblio- 
graphic insight, Adrian Pont (Oxford Uni- 
versity) for deciphering the handwritten la- 
bel of J. Bigot, and Mark A. Metz for 
translating German primary literature. In 
addition, we wish to thank the following 
curators for loans of type materials and for 
their help during the travels of SDG study- 
ing type materials: John E. Chainey 
(BMNH); Loic Matile (MNHN), Ruth Con- 
treras-Lichtenberg (NHMW), and Vadim F 
Zaitsev and Vitali N. Tanasijtshuk (ZMAS); 
and for loans of specimens (including 
types) we thank the following curators, in 
order by collection: David A. Grimaldi 
(AMNH), Richard W. Baumann (BYUC), 
Paul H. Arnaud and Norman D. Penny 
(CASC), Eric M. Fisher (CDFA), Chen W. 
Young (CMNH), Jeffrey M. Cummings 
(CNCI), E. Richard Hoebeke (CUIC), Don- 
ald S. Chandler and John FEF Burger 
(DENH), Enrique Ramirez (EBCC), Cheryl 
B. Barr, John A. Chemsak, and Jerry Pow- 
ell (EMEC), Wilford J. Hanson (EMUS), 
Howard V. Weems, Jr. and Gary J. Steck 
(FSCA), Vicente Hernandez-Ortiz (IEXA), 
Donald W. Webb (INHS), H. Derrick 
Blocker (KSUC), Brian V. Brown (LACM), 
Philip D. Perkins (MCZC), Michael Ivie 
(MTEC), Don C. Arnold (OSEC), Norman 
E Johnson (OSUC), Raymond J. Pupedis 
(PMNH), David Faulkner (SDMC), Robert 
W. Brooks (SEMC), Wade C. Sherbrooke 
(SWRS), Edward G. Riley (TAMU), Carl 
A. Olson (UAIC), Steven L. Heydon and 
Lynn S. Kimsey (UCDC), Saul I. Frommer 
(UCRC), Cecil L. Smith (UGCA), Atilano 
Contreras-Ramos and Harry Brailovsky 
(UNAM), FE Christian Thompson (USNM), 
Frank W. Merickel (WFBM), Richard S. 


599 


Zack (WSUC), H. Wendt (ZMHB), and 
Leif Lyneborg and Verner Michelsen 
(ZMUC). The distributional maps were 
generated using VERSAMAP, version 2.04, 
by Charles H. Culberson. We also extend 
our thanks to FE Christian Thompson, for his 
critical review of the manuscript, and help- 
ful suggestions. Special thanks are due to 
our wives, Helen G. Gaimari and Bonnie J. 
Irwin, for their unending patience and un- 
derstanding. This study was supported by 
NSF DEB-PEET grant # 95-21925 to MEI, 
in addition to funding provided from the 
Schlinger Foundation and a graduate re- 
search assistantship to SDG through the I- 
linois Natural History Survey. 


LITERATURE CITED 


Arnett, R. H., Jr., G. A. Samuelson, and G. M. Nishida. 
1993. The Insect and Spider Collections of the 
World, 2nd edition. Flora and Fauna Handbook 
No. 11. Sandhill Crane Press: Gainesville, 310 pp. 

Becker, T. 1912. Beitrag zur Kenntnis der Thereviden. 
Abhandlungen der Kaiserlich-koniglichen Zoolo- 
gisch-Botanischen Gesellschaft in Wien 62: 289— 
319) 

Bigot, J. M. E 1890. Dipteres nouveaux ou peu con- 
nus, XLIV: Therevidi. Annales de la Société En- 
tomologique de France (1889), Série 6, Vol. 9: 
321-328. 

Bremer, K. 1988. The limits of amino-acid sequence 
data in angiosperm phylogenetic reconstruction. 
Evolution 42: 795-803. 

. 1994. Branch support and tree stability. Cla- 
distics 10: 295-304. 

Carpenter, J. M. 1988. Choosing among equally par- 
simonious cladograms. Cladistics 4: 291—296. 

. 1994. Successive weighting, reliability and 
evidence. Cladistics 10: 215-220. 

Cole, EF R. 1923. A revision of the North American 
two-winged flies of the family Therevidae. Pro- 
ceedings of the United States National Museum 
62(4), 140 pp. 

. 1965. Family Therevidae, pp. 348-354. 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. U.S. De- 
partment of Agriculture Agricultural Handbook 
No. 276. Washington, DC. 1696 pp. 

Donoghue, M. J., R. G. Olmstead, J. E Smith, and J. 
D. Palmer. 1992. Phylogenetic relationships of 
Dipsacales based on rbcL sequences. Annals of 
the Missouri Botanical Garden 79: 333-345. 

Eriksson, T. 1997. AUTODECAY version 2.9.9 (Hy- 


600 


percard stack). Distributed by author: Botaniska 
institutionen, Stockholm University, Sweden. 

Farris, J. S. 1969. A successive approximations ap- 
proach to character weighting. Systematic Zoolo- 
gy 18: 374-385. 

. 1982. Outgroups and parsimony. Systematic 

Zoology 31: 328-334. 

. 1983. The logical basis of phylogenetic anal- 

ysis, pp. 7—36. Jn Platnick, N. L. and V. A. Funk, 

eds., Advances in Cladistics. Proceedings of the 

Willi Hennig Society, Volume 2. Columbia Uni- 

versity Press, New York, 218 pp. 

. 1989. The retention index and the rescaled 
consistency index. Cladistics 5: 417—419. 

Gaimari, S. D. 1998. Phylogeny, classification, and 
biogeography of the cycloteline Therevinae (Dip- 
tera: Therevidae), Ph.D. Dissertation, University 
of Illinois, Urbana. 273 pp. 

Gaimari, S. D. and M. E. Irwin. 2000. Phylogeny, clas- 
sification, and biogeography of the cycloteline 
Therevinae (Diptera: Therevidae). Zoological 
Journal of the Linnean Society (London) 129: 
129-240. 

Godman, FE D. 1901. List of species recorded from 
Mexico or Central America since 1887, not enu- 
merated in the Supplement, pp. 377-378. In God- 
man, FE D. and O. Salvin, eds., Biologia Centrali- 
Americana. Class Insecta. Order Diptera. Volume 
1 (1886-1901). Taylor and Francis: London. 378 
PP- 

Hennig, W. 1966. Phylogenetic Systematics. Univer- 
sity of Illinois Press, Urbana, 263 pp. 

Irwin, M. E., and L. Lyneborg. 198la. The genera of 
Nearctic Therevidae. [Illinois Natural History Bul- 
letin (1980) 32: 193-277. 

1981b. Therevidae, pp. 513-523. In Mc- 
Alpine, J. E, B. V. Peterson, G. E. Shewell, H. J. 
Teskey, J. R. Vockeroth, and D. M. Wood, coords., 
Manual of Nearctic Diptera, Vol. 1. Research 
Branch, Agriculture Canada. Monograph 27, 674 
Pp- 

Kallersj6, M., J. S. Farris, A. G. Kluge, and C. Bult. 
1992. Skewness and permutation. Cladistics 8: 
275-287. 

Kluge, A. G. and J. S. Farris. 1969. Quantitative phy- 
letics and the evolution of anurans. Systematic 
Zoology 18: 1-32. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Krober, O. 1912. Die Thereviden Nordamerikas. Stet- 
tiner Entomologische Zeitung 73: 209-272. 

. 1913. Diptera Fam. Therevidae. Genera In- 

sectorum. Fascicle 148: 69 pp. (in German). 

. 1929. Neue Beitraége zur Kenntnis der Ther- 
eviden und Tabaniden (Dipt.). Deutsche Entomo- 
logische Zeitschrift 1928: 417—434. 

Loew, H. 1869. Diptera Americae septentrionalis in- 
digena. Centuria Octava. Berliner Entomologische 
Zeitschrift 13: 1-52. (In German.) 

Maddison, W. P., M. J. Donoghue, and D. R. Maddi- 
son. 1984. Outgroup analysis and parsimony. Sys- 
tematic Zoology 33: 83-103. 

Nelson, G. J. 1969. Gill arches and the phylogeny of 
fishes, with notes on the classification of verte- 
brates. Bulletin of the American Museum of Nat- 
ural History 141: 475-552. 

. 1972. Phylogenetic relationships and classifi- 

cation. Systematic Zoology 21: 227-231. 

. 1973. Classification as an expression of phy- 
logenetic relationships. Systematic Zoology 22: 
344-359. 

Nixon, K. C. 1999. WINCLADA, Beta version 0.9.9. 
Published by the author. L.H. Bailey Hortorium, 
Cornell University, Ithaca, New York. Available 
as shareware via <http://www. Cladistics.com/> 

Nixon, K. C. and J. M. Carpenter. 1993. On outgroups. 
Cladistics 9: 413-426. 

Sabrosky, C. W. 1978. A third set of corrections to “A 
Catalog of the Diptera of America North of Mex- 
ico.” Bulletin of the Entomological Society of 
America 24: 143-144 

Sanderson, M. J. and M. J. Donoghue. 1989. Patterns 
of variation in levels of homoplasy. Evolution 43: 
1781-1795. 

Swofford, D. L. 1993. PAUP: Phylogenetic Analysis 
Using Parsimony version 3.1.1. Smithsonian In- 
stitution: Washington, DC. 

Swofford, D. L. and W. P. Maddison. 1987. Recon- 
structing ancestral states under Wagner parsimony. 
Mathematics and Bioscience 87: 199—299. 

Watrous, L. E. and Q. D. Wheeler. 1981. The out- 
group comparison method of character analysis. 
Systematic Zoology 30: 1-11. 

Wiley, E. O. 1981. Phylogenetics: The Theory and 
Practice of Phylogenetic Systematics. Wiley Inter- 
science, New York, 439 pp. 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 601-608 


REVIEW OF THE ASIAN SAWFLY GENUS ANISOARTHRA CAMERON 
(HYMENOPTERA: TENTHREDINIDAE) 


V. VASU, DAviID R. SMITH, AND MALKIAT S. SAINI 


(VV, MSS) Department of Zoology, Punjabi University, Patiala-147 002, India; (DRS) 
Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department 
of Agriculture, % National Museum of Natural History, Washington, D.C. 20560-0168, 
U.S.A. (e-mail: dsmith @sel.barc.usda.gov) 


Abstract.—Anisoarthra Cameron includes three species, A. coerulea Cameron from 
India and Sri Lanka, A. diascoreae (Rohwer), n. comb., from India, and A. birmanica 
(Malaise), n. comb., from Burma and India (new record). The species are described and 
illustrated, and a key is provided. Anisoarthra diascoreae feeds on yam, Dioscorea sp. 


(Dioscoreaceae). 


Key Words: 
oclia 


Species of Anisoarthra are relatively 
large, mostly metallic bluish-black or vio- 
laceous sawflies of the subfamily Blenno- 
campinae. A characteristic, shared with sev- 
eral other genera of the subfamily, is the 
comblike, four- or five-toothed tarsal claws. 
The three species of the genus occur only 
in southern Asia—Burma, India, and Sri 
Lanka. Yam, Dioscorea sp., is a host plant 
for Anisoarthra diascoreae (Rohwer). A 
species in a related genus in the Blenno- 
campinae, Senoclidia purpurata (Smith), is 
known to be a pest of yam in Papua New 
Guinea (Szent-Ivany 1974). Since yam is an 
important food plant in the tropics and is 
distributed by commerce, it is important to 
know and be able to recognize its potential 
pests. 

Here we redescribe the species of Ani- 
soarthra and give a key for their separation. 

The National Museum of Natural Histo- 
ry, Smithsonian Institution, Washington, 
D.C., is designated as USNM; Zoological 
Department, Punjabi University, as ZDPU; 
and Canadian National Collection, Ottawa, 
as CNC. 


India, Sri Lanka, Burma, Dioscorea, Dioscoreaceae, Blennocampinae, Sen- 


Anisoarthra Cameron 


Anisoarthra Cameron 1876: 461. Type spe- 
cies: Anisoarthra coerulea Cameron. 
Designated by Rohwer 1911. 

Senoclia Cameron 1877: 88 (unnecessary 
new name for Anisoarthra Cameron).— 
Rohwer 1921: 105 (new species; key to 
species).—Malaise 1937: 50 (separation 
from Neoclia Malaise; new species from 
Burma; key to species).—Benson 1938: 
367 (in Senocliini).—Malaise 1964: 20 
(Anisoarthra not preoccupied and is cor- 
rect name for the genus). 


Description.—Antenna covered with ap- 
pressed hairs; scape about 2 longer than 
broad, pedicel as long as broad, 3rd and 4th 
segments subequal in length, each a little 
dilated at apices; 5th segment dilated at 
apex and little longer than 4th; 6th segment 
about % length of 5th, dilated at apex; 7th 
segment shorter than 6th; apical two seg- 
ments each slightly shorter than 7th (Fig. 
2). Clypeus smooth, truncate or with slight- 
ly rounded anterior margin; malar space lin- 


602 


ear to less than half diameter of front ocel- 
lus; genal carina absent. Epicnemium ab- 
sent. Mesoscutellum raised, conspicuous, 
smooth and shining. Forewing (Fig. 1) with 
vein M and Rs+M meeting before attaining 
Sc+R; vein 2A+3A furcate or curved up 
at its apex, furcation sometimes faint; 4 cu- 
bital cells. Hindwing (Fig. 1) with one mid- 
dle cell; petiole of anal cell as long as cell. 
Tarsal claws comblike, with 4—5 teeth, if 
with 4 teeth, then acute basal lobe present 
(Figs. 3-5). Male penis valve with long api- 
cal filament (Fig. 12). 

Remarks.—The comblike tarsal claws 
are characteristic of several other genera of 
Blennocampinae, such as Neoclia Malaise 
and Brykella Malaise, but Anisoarthra may 
be separated by the venation of the fore- 
wing (vein M and RS+M meeting before 
attaining Sc+R; vein 2A+3A curved up or 
furcate at its apex) and hindwing (petiole of 
anal cell about half as long as cell), absence 
of a genal carina, and absence of an epic- 
nemium. The long apical filament of the 
male penis valve does not occur in other 
known Asian Blennocampinae and appears 
unique to Anisoarthra. 

Much of the literature pertaining to this 
genus has been under the name Senoclia 
Cameron, an unnecessary replacement 
name for Anisoarthra. Cameron (1876) 
originally included two species, A. coerulea 
and A cyanella from Ceylon. Though the 
type of Anisoarthra cyanella is probably 
lost, strong indications are that it belongs to 
Senoclidea Rohwer and that it is not from 
Ceylon as given in the original description 
(see discussion by Smith 1982). In fact, hid- 
den in a footnote, Cameron (1877) stated 
“Tam told by Mr. FE Smith that the locality 
for Senoclia cyanella (l.c. p. 462) is New 
Gunea, and not Ceylon.’’ Consequently, we 
do not include A. cyanella. Rohwer (1921) 
added two species from India, one of which 
was synonymized with A. coerulea by 
Smith (1982), and gave a key to three spe- 
cies. Malaise (1937) added a species from 
Burma and gave a key to species. Malaise 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


did not examine types and based separation 
of Rohwer’s species on the literature. 

Benson (1938) placed Anisoarthra (men- 
tioned as Senoclia) in the tribe Senocliini 
of the Blennocampinae. The tribe included 
a small group of Austro-Oriental genera 
(Kampongia Malaise, Neoclia Malaise, Ne- 
sotomostethus Rohwer, and Anisoarthra) 
separated from other Blennocampinae by 
the point of fusion between veins M and Ir- 
m of the hindwing which reaches to or very 
close to Rs, vein M of the forewing arising 
from vein Rs+M. Nesotomostethus is sep- 
arated from Anisoarthra by the three- 
toothed tarsal claws with a basal lobe and 
presence of an epicnemium. Kompongia is 
separated by the two-toothed tarsal claws 
with a basal lobe and cell M absent in the 
hindwing. 


KEY TO SPECIES OF ANISOARTHRA 


1. Forewing darkly, uniformly infuscated; legs 
black; tarsal claws with 4 teeth and a basal lobe 
(Fig. 3) A. coerulea Cameron 

— Forewing not completely infuscated, approxi- 

mately basal half hyaline; bases of tibiae or 
most of tibiae and extreme apices of femora 
with a whitish or brownish spot; tarsal claws 
with 4 or 5 teeth (Figs. 4—5) 
. Tarsal claws with 4 distinct teeth and a distinct 
triangular basal lobe (Fig. 4); supraclypeal area 
flat; labrum black; lateral furrows on head par- 
allel; postocellar area broader than long, as 4: 
3; about % or less of bases of tibiae whitish to 
brownish; foretibial spur furcate at apex; male 
hypopygium truncate with slight notch at cen- 
tenes. SEO aca ae A. diascoreae (Rohwer) 

— Tarsal claws with 5 distinct teeth, basal lobe 
indistinct, rounded (Fig. 5); supraclypeal area 
triangularly roundly raised; labrum brownish to 
white; lateral furrows diverging posteriorly; 
postocellar area broader than long, as 3:2; basal 
% or foretibia, midtibia except extreme apex, 
and basal % hind tibia white; foretibial spur 
simple; male hypopygium convex, rounded . . 
bags nee ees ee A. birmanica (Malaise) 


i) 


Anisoarthra coerulea Cameron 
(Figs. 3, 8) 

Anisoarthra coerulea Cameron 1876: 462 
(2, 6).—Smith 1982: 188, figs. 1, 9 13 
(2, 6, Sri Lanka; syn.: bilanga Rohwer). 

Senoclia coerulea: Kirby 1882: 181, pl. 8, 


VOLUME 102, NUMBER 3 


603 


Figs. 1-5. 
3, A. coerulea. 4, A. diascoraea. 5, A. birminaca. 


fig. 21; Dalla Torre 1894: 186; Rohwer 
1921: 105 (in key).—Malaise 1937: 52 
(in key; as caerulea). 
Senoclia bilanga Rohwer 1921: 106 (2, 3). 
Senoclia caerulea var. bilanga: Malaise 
19372 D2. 


Female.—Length, 9.0—-11.0 mm. Bluish 
black with metallic hue. Forewing darkly, 
uniformly infuscated; hindwing lightly in- 
fuscated; veins and stigma dark brown. 

Antennal length shorter than abdomen, 
2X head width. Clypeus with anterior mar- 
gin truncate to slightly rounded; labrum 
broader than long, as 2:1, with deflexed and 
roundly pointed anterior margin; supracly- 
peal area triangularly raised; inner margins 
of eyes converging below with lower inter- 
ocular distance to interocular distance at 
level of front ocellus to eye length as 4:5: 


1, Anisoarthra diascoraea, forewing and hindwing. 2, A. diascoraea, antenna. 3—5, Tarsal claws. 


4; malar space linear; head in lateral view 
with frontal area almost on same level as 
eyes and supraantennal tubercles moderate, 
sloping backwards and confluent with low 
frontal ridges; median fovea prominent, 
ditchlike on anterior half and shallow pos- 
teriorly, reaching median ocellus; postocel- 
lar area convex, broader than long, as 3:2; 
temple lateral to lateral ocellus flat; post-, 
inter-, and circumocellar furrows distinct; 
lateral furrows distinct, deep, parallel and 
abruptly ending well before posterior mar- 
gin of head; head from above narrowing be- 
hind eyes; postocellar, ocello-occipital, ocu- 
lo-ocellar, and oculo-occipital distances 
subequal. Mesoscutellum convex, append- 
age not carinated or grooved; distances be- 
tween cenchri to distance between tegulae 
as 1:5. Tarsal claws comblike with 4 dis- 


604 


tinct teeth and triangular basal lobe (Fig. 3); 
foretibial spur furcate at apex; hind basitar- 
sus longer than following 3 segments com- 
bined, as 7:5; hind tibial spurs subequal in 
length, length of inner tibial spur to apical 
width of hind tibia to outer apical tibial spur 
length as 3:2:2. Lancet similar to Fig. 6, 
with 22 serrulae. Sheath short, in lateral 
view with dorsal margin bent down apically 
(Fig. 8). 

Head with sparse, minute, irregular punc- 
tures, surface shining; mesonotum and me- 
sepisternum with minute, scattered punc- 
tures, surface shining with general oily lus- 
ter; mesoscutellum and appendage impunc- 
tate, surface polished; abdomen impunctate, 
surface subshining. Body covered with 
mixed metallic blue and silvery pubes- 
cence. 

Male.—Length, 9.0—10.0 mm. Similar to 
female. Hypopygium truncate with slight 
notch at center. Genitalia similar to Figs. 
12. 

Types.—The lectotype 2 of A. coerulea, 
designated by Smith 1982, is in The Natural 
History Museum, London, labeled *““Type, 
H.T.”; “B.M. Type Hym. 1.363”; ““B.M. 
Type, Hym. Anisoarthra coerulea (Camer- 
ometsro) > Ceylon’: ~Kby-p: Sai, 215” 

The holotype of Senoclia bilanga, a &, 
is in the USNM, labeled ‘“‘Kollegal, 2,000 
feet (about 606 meters), Coimbatoire, S. In- 
dia, 1-[X-17, Ramakrishna.”’ 

Specimens examined.—INDIA: Tamil 
Nadu, Kollegal (Coimbatoire), 600 m, 
25.9.1917; Kollegal, 2,000 ft., 1-[X-17, Ra- 
makrishna, coll. (d allotype of S. bilanga, 
USNM); Nilgiri Hills, Kallar, 1,250 ft., 
South India, Oct. 1955, PS. Nathan. SRI 
LANKA: North Central Province; North 
Western Province; Central Province (see 
Smith 1982 for records). 

Distribution.—India (Tamil Nadu); Sri 
Lanka. 

Host.—Unknown. 

Remarks.—The uniformly infuscated 
wings, entirely bluish black legs, short an- 
tennal length (2 times the head width), and 
4-toothed tarsal claws with a basal lobe will 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


separate this species from both other spe- 
cies of Anisoarthra. The female lancet and 
male genitalia are ver similar to those illus- 
trated for S. diascoreae (Figs. 6, 11-12). 

Rohwer (1921) distinguished bilanga be- 
cause of differences he observed in the 
male. The females he had were identical to 
A. coerulea. The characters he used were 
the uniformly infuscate wings and the head 
more distinctly punctured and with stronger 
antennal furrows. We do not see these dif- 
ferences in the males examined. 


Anisoarthra diascoreae (Rohwer), new 
combination 
(Figs. ly, 24,6; 95 1l=12) 


Senoclia diascoreae Rohwer 1921: 105 (f, 
m).—Malaise 1937: 51 (in key). 


Female.—Length, 10.0—11.5 mm. Bluish 
black with metallic hue; basal ¥% or less of 
tibiae and small spot on apices of femora 
brownish to whitish. Wings darkly infus- 
cated beyond proximal end of cell 1M in 
forewing and hindwing; rest of wings hya- 
line; veins and stigma dark brown to black. 

Antennal length equal to abdomen 
length, subincrassinate in middle, 2.4 
head width; clypeus truncate; labrum broad- 
er than long, as 2:1, with deflexed roundly 
pointed anterior margin; supraclypeal area 
almost flat; inner margins of eyes converg- 
ing below, lower interocular distance to in- 
terocular distance at front ocellus to eye 
length as 7:9:8; malar space about half di- 
ameter of front ocellus; head in lateral view 
with frontal area though prominently raised 
still slightly below level of eyes and su- 
praantennal tubercles slightly indicated and 
confluent with roundly raised frontal ridges; 
median fovea prominent, ditchlike on an- 
terior half and shallow posteriorly, reaching 
median ocellus; postocellar area convex, 
broader than long, as 4:3; temples lateral to 
lateral ocelli depressed, post-, inter-, and 
circumocellar furrows distinct; lateral fur- 
rows distinct, deep, parallel and abruptly 
ending well before posterior margin of 
head; head from above parallel behind eyes; 


VOLUME 102, NUMBER 3 


Figs. 6-12. 6, Lancet of Anisoarthra diascoreae. 7, Lancet of A. birmanica. 8, Female sawsheath of A. 
coerulea. 9, Female sawsheath of A. diascoreae. 10, Female sawsheath of A. birmanica. 11, Ventral view of 
genital capsule of A. diascoraea. 12, Lateral view of penis valve of A. diascoraea. 


606 


postocellar, ocello-occipital, oculo-ocellar, 
and oculo-occipital distances as 4:7:7:5. 
Mesoscutellum subconvex with faintly in- 
dicated lateral carina; mesoscutellar ap- 
pendage neither carinated or grooved; dis- 
tance between cenchri to distance between 
tegulae as 2:9. Tarsal claws with 4 distinct 
teeth and triangular basal lobe (Fig. 4); for- 
etibial spur furcate at apex; hind basitarsus 
longer than 3 following segments com- 
bined, as 5:4; length of inner tibial spur to 
apical width of hind tibia to outer apical 
tibial spur length as 5:3:4. Lancet as in Fig. 
6, with 22 serrulae. Sheath in lateral view 
long wit dorsal margin slightly curved up- 
ward (Fig. 9). 

Head with shallow and scattered punc- 
tures, surface shining, mesonotum with mi- 
nute, scattered punctures, surface shining; 
mesoscutellum with conspicuous punctures 
on posterior slope, surface polished; mesos- 
cutellar appendage impunctate, polished; 
mesepisternun and mesosternum punctured 
like mesonotum, surface shining with gen- 
eral oily luster; abdomen shining with some 
scattered and minute punctures. Body cov- 
ered with mixed metallic blue and silvery 
pubescence. 

Male.—Length, 9.0—10.0 mm. Similar to 
female except antennal segments 7—9 and 
extreme apices of femora brownish. Malar 
space linear; postocellar area broader than 
long, as 3:2. Hypopygium truncate, with 
shallow notch at center. Genitalia as in Figs. 
11-12. 

Types.—The holotype 2 is in the 
USNM, labeled “‘N. Malabar, Taliparamba, 
India’; “‘on pepper vine June ‘18.’ Allo- 
type and paratypes in USNM; paratypes in 
the Zoological Survey of India, Calcutta. 

Specimens examined.—INDIA: Kerala, 
Poonnudi Range (Trivandrum), 3,000 ft, 
May 1972; South Canara Dt., Kollar Ghat, 
3000 ft., 18-21-IX-18, T.V.R. (1 ¢ paratype, 
USNM); N. Malabar, Taliparamba, July— 
Aug. 1918, PS. Nathan (¢ allotype, 
USNM), same data, 16-26-[X-18, Ramak- 
rishna, coll. (1 6 paratype, USNM); Am- 
matti, Coorg, S. India, May 1951, BS. Na- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


than; Nilgiri Hills, S. India, 3,500 ft., May 
1950, P.S. Nathan. 

Distribution.—India (Kerala). 

Host.—Rohwer (1921) stated adults were 
bred ‘‘on a creeper, Diascorea,’’ probably a 
misspelling for Dioscorea sp. (Dioscorea- 
ceae). An adult was collected “‘on pepper 
vine”’ (? black pepper, Piper nigrum L. [Pi- 
peraceae]). 

Remarks.—The tarsal claws with 4 teeth 
and a distinct triangular basal lobe, whitish 
to brownish bases of tibiae and extreme api- 
ces of femora, hyaline basal half of fore- 
wing, furcate foretibial spur, postocellar 
area broader than long, and antennal length 
2.4 times head width, will separate this spe- 
cies from other species of Anisoarthra. 


Anisoarthra birmanica (Malaise), new 
combination 
(Figs..3,; 7710) 


Senoclia birmanica Malaise 1937: 51 (f). 


Female.—Length, 12.0-13.0 mm. Blu- 
ish-black with metallic hue antenna, la- 
brum, extreme apex of forefemur, basal half 
of foretibia, midtibia except extreme apex, 
and basal ¥ of hind tibia whitish to light 
brownish. Forewing and hindwing deeply 
infuscated beyond proximal end of cell 1 M 
and infuscation linearly extending to base 
of wings covering entire anal cell; veins and 
stigma dark brown to black. 

Antennal length equal to abdomen 
length; subincrassinate in middle, 3.2 
head width. Clypeus truncate; labrum 
broader than long as 2:1 deflexed and 
roundly pointed anteriorly; supraclypeal 
area subtriangularly roundly raised; inner 
margins of eyes converging below, lower 
interocular distance to interocular distance 
at level of front ocellus to eye length as 3:4: 
3; malar space linear; head in lateral view 
with frontal area almost at level of eyes and 
supraantennal tubercles significant with 
steep posterior slope and meeting low lying 
frontal ridges; median fovea prominent on 
anterior half and shallow posteriorly, reach- 
ing median ocellus; frontal area anterior to 


VOLUME 102, NUMBER 3 


median ocellus conspicuously depressed, 
thus forming humplike raised projection be- 
tween depression and median fovea; pos- 
tocellar area convex, broader than long, as 
3:2; temples lateral to lateral ocelli subcon- 
vex; post-, inter-, and circumocellar furrows 
distinct; lateral furrows quite distinct, deep, 
diverging backwards and abruptly ending 
well before hind margin of head; head from 
above parallel behind eyes; postocellar, 
ocello-occipital, oculo-ocellar, and oculo- 
occipital distances as 4:4:5:4. Mesoscutel- 
lum subconvex, its appendage neither cari- 
nated nor grooved; distance between cen- 
chri to distance between tegulae as 1:5; tar- 
sal claws comblike with 5 teeth and 
indistinct rounded basal lobe (Fig. 5); for- 
etibial spur simple, hind basitarsus longer 
than following 3 segments combined, as 5: 
4; length of inner tibial spur to apical width 
of hindtibia to outer apical tibial spur length 
as 5:3:4. Lancet as in Fig. 7, with 21 ser- 
rulae; sheath in lateral view long with dor- 
sal margin slightly curved up apically (Fig. 
10). 

Head with dense, irregularly spaced, 
prominent punctures and surface between 
punctures microsculptured, surface shining; 
mesonotum with dense, irregular, minute 
punctures, surface shining; mesoscutellum 
with distinct, irregular, scattered punctures, 
more conspicuous on its posterior slope, 
surface polished; mesoscutellar appendage 
impunctate, polished; mesepisternum and 
mesosternum with few, scattered micro- 
punctures, surface shining with general oily 
luster; abdomen impunctate, shining. Body 
covered with mixed blackish and metallic 
blue pubescence. 

Male.—Length, 10.0—11.0 mm. Similar 
to female. Hypopygium truncate at apex. 
Genitalia similar to Figs. 11—12. 

Types.—Described from 2 ¢, deposited 
in the Naturhistoriska Riksmuseet, Stock- 
holm, Sweden. From Burma (Taungdo at 
the south end of Inle Lake in southern Shan 
States, alt. 900 m, 19.I1X.1934) (Malaise 
1937). 

Specimens examined.—INDIA: Assam, 


607 


Shillong, 1,450 m, 7.6.1943 (ZDPU; As- 
sam, Kameng, Bokhar, 28-V-61, 2,500’ 
(CNC). 

Distribution.—Burma; India (Megha- 
laya). 

Host.—Unknown. 

Remarks.—The mostly whitish to 
brownish tibiae, whitish extreme apices of 
the femora, hyaline basal half of forewing 
and hindwing, triangularly raised supracly- 
peal area, 5-toothed tarsal claws with and 
indistinct rounded basal lobe, white labrum, 
and simple foretibial spurs will separate this 
species from other species of Anisoarthra. 
We did not examine the types. Malaise’s 
(1937) description is sufficient to character- 
ize this species. A series of three females 
and three males from Kameng, Assam as- 
sociate the sexes. 


ACKNOWLEDGMENTS 


Financial assistance rendered by USS. 
PL-480 in collaboration with ICAR is ac- 
knowledged with thanks. We are grateful to 
Henri Goulet, Agriculture and Agri-Food 
Canada, Ottawa (Canadian National Collec- 
tion), for allowing examination of speci- 
mens under his care and for reviewing the 
manuscript. We also thank the following for 
review of the manuscript: A. S. Jensen and 
E. E. Grissell, Systematic Entomology Lab- 
oratory, USDA, Beltsville, MD, and Wash- 
ington, DC, respectively. Cathy Anderson, 
Systematic Entomology Laboratory, helped 
arrange the plates. 


LITERATURE CITED 


Benson, R. B. 1938. On the classification of sawflies 
(Hymenoptera: Symphyta). Transactions of the 
Royal Entomological Society of London 87: 353-— 
384. 

Cameron, P. 1876. VIII. Description of new genera and 
species of Tenthredinidae and Siricidae chiefly 
from the East India, in the collection of the British 
Museum. Transactions of the Entomological So- 
ciety of London, pp. 459-471. 

. 1877. VII. Descriptions of new genera and 
species of East Indian Tenthredinidae. Transac- 
tions of the Entomological Society of London, pp. 
87-92. 

Dalla Torre, C. G. 1894. Catalogus Hymenopterorum, 


608 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Vol. 1. Tenthredinidae incl. Uroceridae (Phyllo- 
phaga & Xylophaga). Lipsiae, 459 pp. 

Kirby, W.E 1882. List of Hymenoptera in the British 
Museum, Vol. 1. London, 450 pp. 

Malaise, R. 1937. New Tenthredinidae mainly from the 
Paris Museum. Revue Frangaise d’Entomologie 4: 
43-53. 

. 1964. New genera and species of the subfam- 
ily Blennocampinae (Hymenoptera, Tenthredini- 
dae). Entomologisk Tidskrift 85: 20-39. 

Rohwer, S. A. 1911. Technical papers on miscella- 
neous forest insects II. The genotypes of the saw- 


flies or woodwasps of the superfamily Tenthredi- 

noidea. United States Bureau of Entomology 

Technical Bulletin 20, pp. 69-109. 

. 1921. Notes on sawflies, with descriptions of 
new genera and species. Proceedings of the Unit- 
ed States National Museum 59: 83-109. 

Smith, D. R. 1982. Symphyta (Hymenoptera) of Sri 
Lanka. Proceedings of the Entomological Society 
of Washington 85: 117—127. 

Szent-Ivany, J. J. H. 1974. Insect pests of yam (Dios- 
corea ssp.) in Papua New Guinea. Folia Ento- 
mologica Hungarica (Series Nova) 28: 205-210. 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 609-612 


CHESPIRITOS, A NEW GENUS OF LIMOSININAE 
(DIPTERA: SPHAEROCERIDAE) FROM COSTA RICA 


S. A. MARSHALL 


Department of Environmental Biology, University of Guelph, Guelph, Ontario, Canada 
N1G 2W1 (e-mail: smarshall @evbhort.uoguelph.ca) 


Abstract.—Chespiritos sindecimus new genus, new species, is described from speci- 
mens taken in decaying bromeliads in Costa Rica. The relationship of Chespiritos to other 


Limosininae is discussed. 


Key Words: 


Chespiritos sindecimus new genus, new 
species, was collected around the leaf-bases 
of wind-downed bromeliads at 2,400 m in 
the Rio Macho reserve, Costa Rica. Most 
of the bromeliads examined were found to 
contain specimens of this new species, as 
well as specimens of an unidentified Prer- 
ogramma Spuletr. 

Terminology in this paper follows Mar- 
shall and Langstaff (1998). Terms used for 
the same structures in other recent papers 
on Sphaeroceridae are indicated in paren- 
theses. 


Chespiritos Marshall, new genus 


Type species.—Chespiritos sindecimus, 
new species. 

Diagnostic characters and similar gen- 
era.—Chespiritos stands out as a distinct 
genus because of its broad head with a large 
and exposed clypeus, two pairs of large in- 
terfrontal bristles, six scutellar bristles, dis- 
tinctive male genitalia, and the complete 
lack of a tenth tergite in the female. The 
most closely related genus is probably 
Sclerocoelus Marshall, with the broad lu- 
nule and well-developed intra-alar bristles 
providing some evidence for this relation- 
ship. Chespiritos, however, lacks the elab- 
orately sclerotized male genital pouch, 


Diptera, Sphaeroceridae, Costa Rica, taxonomy 


complex subcercus, and broad alula which 
define Sclerocoelus (Marshall 1995), and 
Sclerocoelus lacks the basal scutellar bris- 
tles, two interfrontal bristles, highly derived 
phallus, and totally reduced female tergite 
10 (epiproct of authors) which appear to de- 
fine Chespiritos. Similar small basal scutel- 
lar bristles do appear elsewhere in the 
Sphaeroceridae (some species of the Pulli- 
mosina subgenus Dahlimosina Rohacek, 
and three species of Spelobia Spuler, for ex- 
ample). The female tergite 10 of other Li- 
mosininae is occasionally reduced (as in 
some Sclerocoelus), and is occasionally ab- 
sent (as in some Pterogramma with greatly 
reduced cerci), but the complete absence of 
tergite 10 in combination with the presence 
of well-developed cerci is a distinctive at- 
tribute of Chespiritos. 


Chespiritos sindecimus Marshall, 
new species 
(Figs. 1-8) 


Description.—Length from base of an- 
tenna to wing tip 3.0 mm, general color 
brown, tibiae and tarsi luteous. Head strik- 
ingly broad, frontal width between eyes 3X 
height of interfrontal area; clypeus large, 
dark and exposed. Frons with two equal in- 


610 


? Surstylus 


l 


Sternite 5 


Sternite 8 


Sternite 7 7 


Figs. 1-8. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Cercus 
Tergite 8 6 


Tergite 7 


Chespiritos sindecimus. 1-3 and 5, Male abdominal structures. 1, terminalia, left lateral. 2, ter- 


minalia, posterior. 3, aedeagus and associated structures. 5, sternites 5-7. 4, Right wing of male. 6-8. Female 
abdominal structures. 6, tergites 7-8 and cerci. 7, spermathecae. 8, sternites 7—10 and cerci. 


terfrontal bristles and a small lower setula; 
two large orbital bristles; inner occipital 
bristle large, postvertical and postocellar 
bristles very small. Orbit, interfrontal stripe 
and ocellar triangle with indistinct silver 
pollinosity. Palpus clavate, setulose, with 


only weak apical hairs and a single preap- 
ical ventral bristle. First flagellomere flat- 
tened laterally, rounded apically; arista aris- 
ing dorsolaterally, length twice head height, 
aristal hairs long (5X aristal width at mid- 
length). Prosternum bare. Thorax heavily 


VOLUME 102, NUMBER 3 


pollinose; 2 strong postsutural dorsocentral 
bristles separated by 6 rows of acrostichal 
setae; outer postpronotal bristle strong, both 
presutural and postsutural intra-alar bristles 
strong. Scutellum long, with 2 pairs of long 
marginal bristles and a pair of small basal 
bristles. Dorsal surface of mid tibia with 5 
bristles proximally (3 anterodorsal and 2 
posterodorsal; the posterodorsal bristles 
small and sometimes difficult to distinguish 
from posterodorsal setulae) and 5 bristles 
distally (2 anterodorsal, 2 dorsal, 1 poster- 
odorsal). Male mid tibia with an apicoven- 
tral bristle, an anteroventral bristle near 
middle, and a double row of short stout 
ventral bristles on distal half, base of mid 
femur with corresponding stout bristles; fe- 
male mid tibia with a mid ventral and an 
apical ventral bristle. Hind tibia with a 
small anteroventral bristle at apex. Wing 
(Fig. 4) long, third costal sector 0.8 length 
of second, vein R,,; gently sinuate; costa 
extending very far (at least 10 vein-widths) 
beyond apex of vein R,,,;; distance between 
crossveins dm-cu and r-m 3% as long as 
dm-cu, both M,,, and CuA, extending be- 
yond discal cell as pigmented processes; al- 
ula narrow. 

Female abdomen: Tergite 7 with a pale 
posteromedial notch; tergite 8 pale but con- 
vex posteromedially, laterally expanded and 
with differentiated posteroventral parts; ter- 
gite 10 absent (Fig. 6). Cercus short, strong- 
ly tapered, entirely setulose, with 3 small 
outer bristles, a long straight apical bristle 
and a curved inner preapical bristle. Ster- 
nite 7 with a pale posteromedial notch; ster- 
nite 8 reduced to a small, medially pale, 
transverse sclerite with two stout bristles on 
each half; sternite 10 reduced to two small 
plates, one under each cercus (Fig. 7). Area 
between sternite 8 and sternite 10 with a 
large, hyaline vaginal sclerite with a prom- 
inent posteromedial process. Spermathecae 
(3) large, spherical, transversely wrinkled, 
with long conical necks and short scleroti- 
sed parts of ducts (Fig. 8). Pleural mem- 
brane wrinkled and densely setose. 

Male abdomen: Syntergite 1+2 twice as 


611 


long as tergite 3, middle part pale almost to 
hind margin. Sternite 5 small but complex, 
with two pairs of prominent posterior lobes, 
outer pair long-setose and inner pair bare 
and blade-like. Middle part of sternite 5 
dark, very short, with a long deflexed (bent 
upwards) posteromedial lobe ending in two 
small, bristle-bearing lobes (Fig. 5). Ster- 
nite 6 with a narrow ventral part anterior to 
a distinct (but medially membranous) gen- 
ital pouch; a distinct ring sclerite in right 
membrane beside genital pouch. Epandrium 
uniformly long-setose, subanal plate broad, 
weakly bilobed ventrally, subepandrial 
sclerite broad and arching slightly above 
level of subanal plate; each half of subanal 
plate with a single long bristle (Fig. 2). Sur- 
stylus leaf-like, with a row of stout bristles 
on anterior edge and a basal patch of fine 
bristles (Fig. 1). Hypandrium stout, deeply 
cleft posteriorly and with an elongate apo- 
deme anteriorly, anterior arms long and 
fused both with apodeme and epandrium, 
no median posterior part but with small 
lobe connecting anterior arm to postgonite, 
pregonite (suspensory sclerite) small and 
closely appressed to postgonite. Postgonite 
(paramere, gonostylus) dark, flattened, apex 
serrate with a posteroapical lobe (Fig. 3). 
Basiphallus stout, wedge-like; distiphallus 
heavily sclerotised, tubular basally and with 
two detached distal dorsal lobes and a spat- 
ulate ventral lobe. 

Type material.—Holotype (d¢, INBio) 
and 9 paratypes (2 2, 2 d, INBio; 2 2, 3 
6 GUELPH): COSTA RICA. Cartago, Rio 
Macho, La Esperanza, El Guarco (8 km 
from km 61 Interamerican Highway) 2,400 
m, LN-188200, 549800, Hoja Tapanti, in 
bromeliads on ground, 1.vi.1998, S. A. 
Marshall. Holotype and two paratype @ 
with two large mites each. 

Etymology.—The genus is named after a 
well-known truck stop on the Interamerican 
Highway near the type locality; the gender 
is masculine. The specific name refers to 
the lack of tergite ten on the female abdo- 
men. 

Comments.—The type series of Chespir- 


612 


itos sindecimus was collected as part of an 
INBio (Instituto Nacional de Biodiversidad, 
Costa Rica) sponsored field trip for partic- 
ipants in a planning workshop for a major 
biodiversity inventory project in Costa 
Rica, and this description can be viewed as 
an initial contribution to that inventory. 


ACKNOWLEDGMENTS 


Thanks to INBio (Instituto Nacional de 
Biodiversidad, Costa Rica), and especially 
dipterist Manuel Zumbado, for making it 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


possible for me to collect this and many 
other interesting Sphaeroceridae in Costa 
Rica. 


LITERATURE CITED 


Marshall, S. A. 1995. Sclerocoelus and Druciatus, new 
genera of New World Sphaeroceridae (Diptera: 
Sphaeroceridae: Limosininae). Insecta Mundi 9: 
283-289. 

Marshall, S. A. and R. Langstaff. 1998. A revision of 
the New World Opacifrons Duda (Diptera, 
Sphaeroceridae, Limosininae). Contributions in 
Science 474: 1-27. 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 613-624 


A REVIEW OF THE FLEA GENERA HECTOPSYLLA FRAUENFELD AND 
RHYNCHOPSYLLUS HALLER (SIPHONAPTERA: PULICIDAE) 


MICHAEL W. HASTRITER AND EUSTORGIO MENDEZ 


(MWH) Monte L. Bean Life Science Museum, Brigham Young University, 290 MLBM, 
P.O. Box 20200, Provo, Utah 84602-0200, U.S.A. (e-mail: hastritermw @ sprintmail.com); 
(EM) Instituto Conmemorativo Gorgas de Estudios de la Salud, Apartado Postal 6991, 


Zona 5, Panama, Republica de Panama 


Abstract.—Rhynchopsyllus, a monotypic genus, is proposed as a new junior synonym 
of Hectopsylla. A key for the 12 species of Hectopsylla is provided with annotation of 
host preferences and geographical distribution of Hectopsylla pulex (Haller). 


Key Words: 


Rhynchopsyllus pulex described by Hall- 
er (1880) has been a source of taxonomic 
confusion. Schreiter and Shannon (1927) 
and Traub and Gammons (1950) erected 
Maxilliopsylla lilloi and Rhynchopsyllus 
megastigmata, respectively, from females 
only. Tipton and Mendez (1966) subse- 
quently described the male of R. megastig- 
mata from Panama (in the presence of ac- 
companying females). Later Méndez (1977) 
reported associated males and females from 
Colombia to be the same as R. megastig- 
mata from Panama, but considered them 
conspecific with R. pulex. Jordan (1939) 
synonymized M. lilloi with R. pulex. Jordan 
and Rothschild (1906), Dalla Torre (1924), 
and Pinto (1930) each reported the species 
as Hectopsylla (based only on females). In 
addition, Anduze et al. (1947) and Cova 
Garcia and Tallaferro (1959) questioned the 
status of Rhynchopsyllus because of simi- 
larities with Hectopsylla. If one uses the 
characters described by Hopkins and Roths- 
child (1953), Panamanian and Colombian 
males reported by Tipton and Méndez 
(1966) and Méndez (1977) clearly belong 
to Hectopsylla. This paper provides obser- 
vations to support synonymizing the mono- 


Hectopsylla, Rhynchopsyllus, Siphonaptera, Pulicidae, key 


typic genus Rhynchopsyllus with Hectop- 
sylla. 

Mammalian synonymies follow those of 
Wilson and Reeder (1993) and avian syn- 
onymies are those described in Peters 
(1934, 1940), Deignan et al. (1964) and 
Sibley and Monroe (1990). The deposito- 
ries of material examined for this study are 
annotated after host/locality data as: The 
Natural History Museum, London 
(BMNH), Field Muséum of Natural Histo- 
ry, Chicago (FMNH), J.C. Beaucournu Col- 
lection (JCB), Museum d’ Histoire Naturel- 
le, Belgium (Md’HN), Michael W. Hastriter 
Collection (MWH), Robert E. Lewis Col- 
lection (REL), and National Museum of 
Natural History, Smithsonian Institution, 
Washington D.C. (USNM). 


Genus Hectopsylla Frauenfeld 


Hectopsylla Frauenfeld 1860: 464. Type 
species (by monotypy): Hectopsylla psit- 
taci Frauenfeld. 

Rhynchopsyllus Haller 1880: 72. Type spe- 
cies (by monotypy): Rhynchopsyllus pu- 
lex Haller. New synonymy. 

Maxilliopsylla Schreiter and Shannon 1927: 
6. Type species (by monotypy): Maxil- 


614 


liopsylla lilloi Schreiter and Shannon. 
Synoymized by Jordan 1939: 303. 


The diagnostic features distinguishing 
Rhynchopsyllus from Hectopsylla have 
been based only on females of the former. 
Jordan (1934) and Hopkins and Rothschild 
(1953) based separation of these two genera 
by the morphology of the spermatheca and 
maxilla. Jordan (1934) specifies the differ- 
ences as ‘a long, narrow, pointed and 
curved maxilla and a conical projection at 
the orifice of the spermatheca”’ in Rhyn- 
chopsyllus, whereas the maxilla of Hectop- 
sylla is ‘“‘short and broadly triangular and 
the orifice of the spermatheca is flush with 
the surface, not projecting on a cone”’. 
More recent descriptions of the male of R. 
pulex by Tipton and Méndez (1966) and 
two additional species (H. gracilis Mahnert, 
1982 and H. pascuali Beaucournu and A\l- 
cover, 1989) provide evidence that would 
suggest Rhynchopsyllus is synonymous 
with Hectopsylla. Hectopsylla gracilis and 
H. pascuali also have sharply pointed max- 
illae that are directed caudad. Both species 
clearly belong to Hectopsylla. Comparison 
of these three species illustrates the simi- 
larity of their maxillae (Figs. 19-21). Rhyn- 
chopsyllus pulex is the only species among 
the two genera, which bears a broad conical 
projection at the orifice of the duct of the 
spermatheca. Although the spermatheca is 
morphologically distinct from that of all 
species of Hectopsylla (except H. stomis to 
which it is quite similar) the authors con- 
sider the conical structure relevant as a spe- 
cies distinction only. 

Both genera share a common sessile par- 
asitic mode of life (primarily in the fe- 
males). Modifications for a sessile mode of 
life shared by females of both genera in- 
clude extreme serration and sometimes ex- 
treme elongation of the lacinia, angular 
frons, compression of thoracic segments, 
partial covering of the anterior portion of 
antennal fossa, and expansibility of inter- 
segmental abdominal membranes (not true- 
ly neosomic). The expansibility of interseg- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


mental abdominal membranes is demon- 
strated most dramatically among females of 
R. pulex, H. knighti, and H. psittaci (the 
only species within the two genera with 
preference for volant hosts). This is most 
extreme in R. pulex and is accompanied by 
autoseverence of appendages (a feature of 
species of Tunga). Evidence of autosever- 
ence in R. pulex includes catabolic scarring 
of remaining coxae and trochanters, absent 
in all species of Hectopsylla examined. 

Neither males of Rhynchopsyllus, nor 
Hectopsylla have developed adaptations to 
accomodate a sessile mode of life (shorter 
mouthparts, fewer serrations on lacinia, ten- 
dency for rounded frons, and inexpansible 
abdomens as demonstrated by examination 
of mounted male specimens of Hectopsylla 
previously attached to hosts). The feeding 
and copulatory behavior of Rhynchopsyllus 
or Hectopsylla males is virtually unknown. 
Their lack of abundance on hosts (none for 
Rhynchopsyllus) would suggest that feeding 
is minimal (if at all) and the occasional col- 
lection of male specimens of Hectopsylla 
from a host likely occurs while they are 
seeking females for mating. 

The clasper of Rhynchopsyllus is closely 
allied with that of Hectopsylla, possessing 
a posteriorly projecting manubrium termed 
manubrium 2 by Hopkins and Rothschild 
(1953). The pattern of the aedeagus of 
Rhynchopsyllus is also similar to those of 
all species of Hectopsylla. The sclerotized 
inner tube (S.I.T.) is nearly identical, being 
extremely long and narrow with a small 
basal capsule to guide the short penis rod 
into the S.I.T. (Figs. 5—6, 8). Dorsal to the 
capsule and running obliquely parallel to 
the S.I.T. is a heavily sclerotized structure 
referred to as the crescent sclerite by Tipton 
and Méndez (1966) in their description of 
R. pulex males. This pair of sclerites is pre- 
sent in all Hectopsylla species, as well as 
R. pulex, but is not present in Tunga, or 
Echidnophaga (the most closely allied gen- 
era). Since the structure is dorsal and dis- 
tinct from the defined capsule of the aedea- 
gus (and not the roof of the capsule = cres- 


VOLUME 102, NUMBER 3 


615 


Table 1. A summary of specimens examined during this study illustrating host preferences within the genus 


Hectopsylla. 

Non-Chiropteran Host 

Flea species Chiropteran Mammal Aves Unknown 

H. broscus 10) 1/13! 0) 0) 
H. coniger 0 1/7 0) 0) 
H. cypha 0 5/74 0 0) 
H. eskeyi 0 5/40 0 10) 
H. gemina 0) 18/54 0) 8/6 
H. gracilis 0) 2/9 0) 0) 
H. knighti 0) 0) 0/1 0) 
H. pascuali 0) 1/2 0) 10) 
H. psittaci O 6) 5/61 0/5 
H. pulex 0/34 0/3 O 3/3 
H. stomis 0) WAS) 0/5 0/4 
H. suarezi 0) 5/6 0) 0/1 


0/0 = number of males/number of females. 


cent sclerite), it is not the crescent sclerite. 
These sclerites are herein referred to as the 
dorsal armature (D.A.) (Figs. 5—6). The 
apex of the D.A. of specimens of all species 
appears fused or at least contiguous with 
the S.I.T. (Fig. 6) (the position of the D.A. 
as illustrated in Fig. 5:is atypical of most 
specimens examined). Based on the com- 
mon morphological features discussed, the 
authors consider the monotypic genus 
Rhynchopsyllus a junior synonym of Hec- 
topsylla. 


Hectopsylla pulex (Haller) 
(Figs. 6—7, 19) 


Rhynchopsyllus pulex Haller 1880: 72. 

Rhynchopsylla_ pulex: Taschenberg 1880: 
56. 

Hectopsylla psittaci Baker 1904: 375, 434. 

Hectopsylla pulex: Jordan and Rothschild 
19065.59;.63. 

Rhynchopsyllus pulex: Cunha 1914: 172. 

Hectopsylla pulex: Dalla Torre 1924: 20. 

Maxilliopsylla lilloi Schreiter and Shannon 
1927: 6 (type species of Maxilliopsylla, 
by monotypy). 

Rhynchopsyllus pulex: Ewing 1929: 158. 

Hectopsylla pulex: Pinto 1930: 332. 

Rhynchopsyllus pulex: Jordan 1934: 19; 
Jordan 1939: 303 (synonymy of M. /il- 
loi); Guimaraes 1940: 219; Fuller 1942: 


44; Anduze et al. 1947: 1-10; Macchia- 
vello 1948: 15; Augustson and Ryan 
1948: 111. 

Rhynchopsyllus megastigmata Traub and 
Gammons 1950: 271; Hopkins and 
Rothschild 1953: 68. 

Rhynchopsyllus pulex: Hopkins and Roths- 
child 1953: 66; Johnson 1957: 237; Bar- 
rera and Diaz-Ungria 1957: 174. 

Hectopsylla pulex: Cova Garcia and Tallaf- 
erro, 1/9592 35. 

Rhynchopsyllus megastigmata: Tipton and 
Méndez 1966: 296. 

Rhynchopsyllus pulex: Tamsitt and Fox 
1970: 1093; Tipton and Machado-Allison 
1972: 4; Méndez 1977: 164 (synonymy 
of R. megastigmata). 

Material examined.—ARGENTINA: 
Buenos Aires, 1 2, Ex: Nyctinomus bras- 
iliensis = Tadarida brasiliensis (1. Geof- 
froy) (BMNH). BOLIVIA: S. Cruz de la 
Sierra, 2 2, Ex: Molossus obscurus = 
Molossus molossus (Pallas) (BMNH); 
Magdalena, Dept. Beni, 2 2, Ex: Eumops 
bonariensis (Peters), 12 Nov 1966; Mag- 
dalena, Dept. Beni, 1 2, Ex: Noctilio la- 
bialis = Noctilio albiventris Desmarest, 
20 Aug 1964 (USNM). BRAZIL: 1 
(paratype), Ex: Molossus sp.; Lagoa San- 
ta, 1 2, Ex: Eumops perotis (Schinz), 2 
Feb 1916 (BMNH); Lagoa Santa, 2 °, 


616 


Ex: E. perotis, 2 Jan 1944 (USNM); Pe- 
tropolis, 1 2, Ex: “‘ear of bat’’; Parana 
Prov., 3 2°, Ex: Histiotus velatus (1. Geof- 
frey) (BMHH). COLUMBIA: Anserman- 
uevo, Dept. Valle, 4 2, Ex: M. molossus, 
20 Jul 1967; Cali, Dept. Valle, 2 2, Ex: 
Molossus major = M. molossus, 22 Nov 
1962; Cali, Dept. Valle, 12, Ex: M. mo- 
lossus, Jun 1967; Cartago, Dept. Valle, 2 
2, Ex: Molossus bondae J.A. Allen, 10 
Oct 1967 (USNM). ECUADOR: Albitig- 
ua, elev. 1,000 m, 3 2, Ex: Rhipidomys 
leucodactylus (Tschudi); Gualquiza, elev. 
760 m, 12, Ex: ‘“‘bat’?. PANAMA: Pa- 
cora, 2d, Ex: “bat guano’’, 22 Jun 1961 
and 27 Feb 1962 (USNM); Pacora, 1 6, 
ie oeExe bat guano’, 20, 22 Jun 1961: 
Pacora, 2 °, Ex: “‘bat guano’’, 12, 14 Feb 
1962; (MWH). PERU: Rio Chinchao, 
Dept. Huanuco, | 2, Ex: Molossus ob- 
scurus = M. molossus (BMNH); Quince 
Mil, Cuzco Prov., 1°, Ex: T. braziliensis, 
19 Jun 1950 (USNM). TRINIDAD: San- 
tissimas il! “2 Exe’ “bat-4-27 Jul i915 
(USNM). UNITED STATES: Frio Cave, 
Uvalde County, Texas, 1 2, Ex: Tadarida 
mexicana = T. brasiliensis, 30 Mar 1955 
(USNM). VENEZUELA: Esteban, 
Djiques, 2 2, Ex: M. obscurus; Merida, 
1 2, Ex: Vespertilio fuscus = Eptesicus 
fuscus (Beauvois) (BMNH); 3 km S and 
46 km W Caracas, Dto. Federal, 2 2, Ex: 
M. obscurus = M. molossus, 19, 20 Aug 
1966 (USNM). 

Remarks.—Hectopsylla pulex, one of 
12 species of Hectopsylla, occurs primar- 
ily on bats of the family Molossidae and 
to a lesser extent on Vespertilionidae. The 
remaining species parasitize either Aves 
(HM. knighti, H. psittaci) or non-Chirop- 
teran mammals (Caviidae: H. cypha, H. 
eskeyi, H. gemini, H. suarezi; Muridae: 
H. pascuali, H. gracilis; Mustelidae: H. 
broscus, H. coniger; and Chinchillidae: 
H. stomis) (Table 1). The known country 
records of Hectopsylla pulex, its host spe- 
cies and host synonymies are listed in Ta- 
ble 2. The occurrence of a single female 
of H. pulex on Zonotrichia pileata = 


Nw 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Zonotrichia capensis (Miller) (reported 
in Cunha 1914) and three females ex- 
amined from the Murid rodent R. leuco- 
dactylus are likely accidental associa- 
tions. 

Little is known about the life cycle of 
H. pulex. Females have been collected 
frequently on chiropteran hosts while 
males have been collected only from bat 
guano associated with species of Molos- 
sus in Colombia and Panama. 


KEY TO SPECIES OF HECTOPSYLLA 


Maxilla rounded at apex (Fig. 11) (male un- 
known) knighti 
Maxilla broadly angular to sharply pointed at 
apex (Bigs: 1222) oie cs eae a ee 2 
Segment V of all tarsi with seven or eight 
(usually eight) lateral planter bristles; tarsal 
claws with a prominent basal tooth; metepi- 
meron of female without a process on dorso- 
posterior margin of metepimeron psittaci 
Segment V of all tarsi with at most six pairs 
of lateral plantar bristles; tarsal claws without 
a prominent basal tooth; metepimeron of fe- 
male nearly always with a process at dorso- 


posterior margin (Fig. 2) (except pulex) ... 3 
Malet. fie afeiiege & 2c A eee 4 
Femalé;..¢ cbs ssieacite a betel th eee 13 


P! of clasper narrower at apex than at base 
(Figs. 3, 7) 
P! of clasper broader at apex than at base (Fig. 
A) verses ak suyex genie hit ee Ee eee 7 
Tarsal segments V with six pairs of lateral 
plantar bristles; median dorsal lobe of aedea- 
gus absent or vestigial ............. broscus 
Tarsal segments V with five or fewer pairs of 
lateral plantar bristles on tarsi; median dorsal 
lobe well developed (Fig. 5) 
P? and P® of clasper pincer-like (Fig. 4); distal 
arm of st. [IX divided into several lobes; fifth 
tarsal segments with four pairs of lateral plan- 
tar"bristles "iss seg. a ee ee © oe stomis 
P? and P® not pincer-like (Fig. 7); distal arm 
of st. [IX composed of a single lobe; fifth tar- 
sal segments with five pairs of lateral plantar 
DTIStleS yas cece meee gaeys ok ee eee pulex 
Median dorsal lobe of aedeagus poorly de- 
veloped, at most discernible 
Median dorsal lobe well developed (Fig. 5), 
distinctly visible 
Median lobe (L?) of distal portion of st. IX 
enlarged with concavity at ventral margin; 
apical margin of P! of clasper straight; P* not 
bifurcate apically gracilis 
Ventral margin of L? lacking concavity; apical 


VOLUME 102, NUMBER 3 


617 


Table 2. Distribution and host-parasite relationships of Hectopsylla pulex. 


Country 


Host species! 


Flea sex 


Argentina 


Myotis nigricans (Schinz) 


2 


Nyctinomus brasiliensis = Tadarida brasiliensis (1. Geot- 2 


froy) 


Promops perotis = Eumops perotis (Schinz) 
Zonotrichia pileata = Zonotrichia capensis? (Miiller) 
Eumops bonariensis (Peters) 


Bolivia 


Molossus obscurus = Molossus molossus (Pallas) 
Noctilio labialis = Noctilio albiventris Desmarest 


Brazil E. perotis 


Histriotus velatus = Histiotus velatus (11. Geoffroy) 
Molossus bondae J. A. Allen 

Molossus obscurus = M. molossus 

Molossus rufus = Molossus ater E. Geoffroy 
Nyctinomus brasiliensis 


Colombia Molossus bondae 


Molossus major = M. molossus 
Molossus molossus major = M. molossus 


M. molossus 


Molossus obscurus obscurus = M. molossus 
Noctilio labialis = Noctilio albiventris 


‘““Bat’? 


Rhipidomys leucodactylus* (Tschudi) 


E. perotis 


Geoffroy) 
Histiotus sp. 


Molossus obscurus = M. molossus 


Tadarida brasiliensis 
Salata 

E. perotis 

Tadarida mexicana = 


Trinidad 
United States 


Venezuela 


T. brasiliensis 
Nyctinomus macrotis = Nyctinomops macrotis (Gray) 


Tadarida yucatanica = Nyctinomops laticaudatus (E. 


T. brasiliensis 
M. major = M. molossus 


M. obscurus = M. molossus 


Myotis nigricans 


Vespertilio fuscus = Eptesicus fuscus (Beauvois) 


+O +0 


Oy Oy 
 40)4+0) 40 FO GO" — 40) +0) +O 40) 40) 4OT4O 407410 40 40 tORtO tO 


Oy 
+O 


+O +O +O +0 +0 +0 +0 +0 +0 +0 


' Host synonymy after Wilson and Reeder (1993). 
? Only known record occurring on a bird. 


3 Only known record occurring on a non-Chiropteran mammal. 


margin of P! somewhat convex; P2 strongly 
bifurcated into two dentate projections 
pascuali 
9. Dorso-posterior margin of metepimeron with 
a caudally directed process (Fig. 2) 
— Dorso-posterior margin of metepimeron en- 
tirewlackinoyprocess oe Sood se. 4s] - 10 
10. Sclerotized inner tube (S.I.T.) with a sharp 
thorn-like spine at ventral midpoint (Fig. 8); 
apex of ventral lobe (L*) of distal portion of 
st. [IX pointed; fifth tarsal segments with three 
pairs of lateral plantar bristles (some with 
three and four on same metatarsus) . gemina 
— §S.LT. without thorn-like spine at ventral mid- 


. coniger 


point, if with tubercle or protuberance then 
rounded and blunt (Fig. 5); apex of L* round- 
ed; fifth tarsal segments with either four or 


five pairs of lateral plantar bristles ...... 11 
. L? with lateral patch of long thin setae; usu- 
ally four pairs of lateral plantar bristles on 
fiithytarsalfscoments ee ae een ie cypha 
L? without lateral patch of setae, but with ver- 
tical line of setae; four or five pairs of lateral 
plantar bristles on fifth tarsal segments .. . 12 
. Fifth tarsal segments with four pairs of lateral 
plantar/bristles® ets. <5. a ee oe ae eskeyi 
Fifth tarsal sements with five pairs of lateral 
platitarsbristles, . "2088 Pee 2 eee suarezi 


618 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


9 10 


Figs. 1-10. 1, Head, female H. broscus. 2, Metepimeron, H. eskeyi (female paratype). 3, P' of clasper, H. 
stomis. 4, Clasper, H. eskeyi (holotype). 5, terminal portion of aedeagus, H. eskeyi (holotype), 6, Terminal prtion 
of aedeagus, H. pulex. 7, Clasper, H. pulex. 8, Sclerotized inner tube, H. gemina. 9, Metatarsal claw, H. pascuali 
(allotype). 10, Metatarsal claw, H. gracilis (female paratype). Abbreviations: CR. = crochet; C.S. = crescent 
sclerite; D.A. = dorsal armature; L.L. = lateral lobe; L.S. = lateral sclerite; M.D.L. = medium dorsal lobe; P', 
P2 and P? = processes of clasper; PS.C. = pseudocrochet; S.I.T. = sclerotized inner tube. Scale = 100p. 


VOLUME 102, NUMBER 3 


13. Metepimeron with a dorso-posterior projec- 
tion (Fig. 2) 
— Metepimeron without a dorso-posterior pro- 
jection pulex 
14. Posterior margin of occiput with a well de- 
finedwlobe (bisa) tees see ce wea ene eee 15 
— Posterior margin of occiput without a well de- 


fined lobe (though slightly indicated in eskeyi) 


15. Tergum II with three setae per side and t. VII 
with two or three setae per side; frons with 
convex angle midway between oral angle and 
internal incrassation (Fig. 1); five or six (usu- 
ally six) lateral plantar bristles on fifth tarsal 
SECMENtS Aaesissloticbs CG.cs ae a Sew broscus 

—  Tergum II with one or two setae per side and 
t. VII with one per side; frons with angular 
projection immediately below internal incras- 
sation; three lateral plantar bristles on fifth 
(RICA SSSNETNIG 5 poo ea pom co ooo oc coniger 

16. Sclerotized rim of sensilial plate anterior to 
sensilial pits greater than twice width of dou- 
blesowsOl pits ef cei Pi sns agls: Sacp tego ee stomis 

—  Sclerotized rim of sensilian plate anterior to 
sensilial pits at most as wide as double row 
of pits 

17. Fifth tarsal segments with three pairs of lat- 
eral plantar bristles; ventral apical margin of 
t. VIII with tooth-like projection 


gemina 

— Fifth tarsal segments with either four or five 
pairs of lateral plantar bristles; ventral apical 
margin of t. VIII without tooth-like projection 


18. Fifth tarsal segments with five pairs of lateral 
plantar bristles (occasional specimens may 
have four and five on the same _ tarsus) 

Pe Neher b ot Picacho oats a aeaee Shad 2-3 ketaars suarezi 

— Fifth tarsal segments with four pairs of lateral 
Plantarbnistlesse ee serene ee ae 19 

19. Dorsal margin of metepimeron heavily scler- 
otized, sclerotization extending to apex of 
process which is markedly turned down; usu- 
ally three setae on metepimeron ....... cypha 

— Dorsal margin of metepimeron not noticeably 
sclerotized (Fig. 2); usually four setae on me- 

He PIMELOME foe eee ob a ea wen meets S 20 

20. Hilla of spermatheca nearly as wide through- 
out as width of bulga; duct of spermatheca 
connecting at cribriform area of bulga on ven- 

Teal MAT OWN wma. tte cis acc fies Sees ioe eskeyi 

— Hilla of spermatheca much narrower than 
width of bulga, with marked narrowing from 
base of bulga to apex of hilla; duct of sper- 
matheca connecting at cribriform area of bul- 


ga on ventral apical margin ........... 21 
21. Base of metatarsal claw with small cleft, or 
CHHAUNS (CS ex OD) A) Gi Stu au 6 Blond ote. 0, Sto oe pascuali 


619 


— Base of metatarsal claw without cleft, or sinus 
(Fig. 10) gracilis 


OTHER MATERIAL EXAMINED 


Hectopsylla broscus Jordan and 
Rothschild 1906 
(Fig. 12) 


Material examined.—ARGENTINA: 
Central Pampa, 1 2? (lectotype), Ex: Co- 
nepatus humboldtii Gray; Central Pampa, 5 
2 (paralectotypes), Ex: C. humboldtii; San 
Rafael, Mendoza Prov., 1 6, 1 2, Ex: Co- 
nepatus suffocans = C. chinga (Molina); 
Don Roberto, San Luis Prov., 1 2, Ex: Co- 
nepatus chinque = C. chinga, 14 Jun 1962; 
‘Southern Argentina,’ 5 2, Ex: Zaedyus 
pichiy (Desmarest), 30 Jun 1962 (BMNH). 

Remarks.—All records of this species are 
restricted to Argentina and skunks of the 
genus Conepatus appear to be the preferred 
host. 


Hectopsylla coniger Jordan and Rothschild 
1906 
(Fig. 17) 


Material examined.—BOLIVA: Pampa 
Olliga, 1 ¢ (lectotype), Ex: Conepatus ar- 
equipae = C. chinga, 19 Oct 1901; Pampa 
Olliga, 1 2 (lectoallotype), 6 2 (paralec- 
totypes), Ex: C. chinga, 19 Oct 1901 
(BMNB). 


Hectopsylla cypha Jordan 1942 
(Fig. 14) 


Material examined.—ARGENTINA: Las 
Catitas, Mendoza Prov., 9 2 (paratypes), 
Ex: Octomys barrerae = Tympanoctomys 
barrerae (Lawrence), Jul 1939; Mendoza, 1 
2, Ex: Microcavia australis (1. Geoffroy 
and d’Orbigny), 6 Aug 1959; Puesto “‘La 
Carpa,’’ Mendoza, 5 2, Ex: Graomys gri- 
seoflavus griseoflavus = Graomys griseo- 
flavus (Waterhouse), 12, 14 Jul 1959; Pues- 
to ‘“‘La Carpa,’”’ Mendoza, 2 2, Ex: M. aus- 
tralis, 13, 15 Jul 1959; San Rafael, Men- 
doza Prov., 1 2 (neoallotype), 2 ¢, 52 @ 
(paratypes), Ex: M. australis, Apr—Jul 
1939; Santa Rosa, Mendoza Prov., 2 6, 4 
2 (paratypes), Ex: M. australis, Jul 1939 


620 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


19 20 21 22 


Figs. 11-22. First segment of maxillary palpus, maxilla, and ventral margin of head (genal lobe); male is 
positioned directly above female for each species. 11, H. knighti (female holotype). 12, H. broscus (female 
lectotype). 13, H. stomis (female holotype). 14, H. cypha (paratypes). 15, H. gemina. 16, H. suarezi. 17, H. 
coniger (lectotype and allolectotype). 18, H. eskeyi (holotype and paratype). 19, H. pulex. 20, H. pascuali 
(holotype and paratype). 21, H. gracilis (holotype and paratype). 22, H. psittaci. Scale = 100w. 


VOLUME 102, NUMBER 3 


(BMNH). Zapallar, La Rioja Prov., 1 ¢, Ex: 
*“Viscacha”’ Lagostomus sp. Brookes, 8 Jan 
1933 (USNM). 

Remarks.—Mountain cavies (M. austral- 
is), which are restricted to Argentina and 
southern Chile, are the preferred host for 
this flea. The geographic distribution of T. 
barrerae is also restricted to Argentina and 
it is rarely collected. Hectopsylla cypha oc- 
curring on other hosts that have much 
broader geographical distributions (Lagos- 
tomus sp. and G. griseoflavus) are likely ac- 
cidental associations. 


Hectopsylla eskeyi Jordan 1933 
(Figs. 2, 4—5, 18) 


Material examined.—BOLIVIA: Pucara, 
elev. 2,400 m, 1 2, Ex: G. griseoflavus, 28 
Sep 1954; Samaipata, Dept. Santa Cruz, 
elev. 1,650 m, 1 ¢, Ex: G. griseoflavus, 28 
Jan 1955; Serrano, Dept. Chuquisaca, elev. 
2,160 m, 1 2, Ex: Hesperomys muriculus 
= Calomys callosus (Rengger), 24 Jul 1955 
(BMNH). ECUADOR: Riobamba, Chim- 
borazo Prov., elev. 2,800 m, 1 @°, Ex: Rat- 
tus rattus (Linn.), 12 Jun 1956; Sanjapam- 
ba, Tungurahua Prov., 6 2, Ex: Cavia por- 
cellus (Linn.), 1956 (BNMH). PERU: 
Huancabamba, elev. 1,960 m, 4 6,9 2, Ex: 
Cavia cobaya (domestic) = C. porcellus, 
17, 19 May 1956; Lima, 1¢ (holotype), 1 
2 (Neoallotype), 1 ¢, 18 @ (paratypes), 
Ex: “rats,” 1930; Yura, Arequipa Prov., 
elev. 8,200 ft, 1 2, Ex: Cavia musteloides 
= Galea musteloides Meyen, 8 Aug 1939 
(BMNH); Yura, Arequipa Prov., elev. 8,200 
ft, 1 2, Ex: G. musteloides (REL). 

Remarks.—Hectopsylla eskeyi occurs in 
the higher Andean valleys on a variety of 
hosts and little can be said of its host spec- 
ificity. 


Hectopsylla gemina Jordan 1939 
(Figs. 8, 15) 


Material examined.—ARGENTINA: 
Mendoza, | ¢, 4 2, Ex: M. australis, 9 Aug 
1959; Las Catitas, Mendoza Prov., 1 3, 1 
2, Ex: Octomys barrerae = T. barrerae; La 
Paz. Mendoza\Prov., 3° 6, 232 ,°Ex: M. 


621 


australis, Feb-Mar 1939; Puesto “‘La Car- 
pa,’ Mendoza Prov., elev. 600 m, 1 6, 6 
2, Ex: M. australis, 13-17 Jul 1959; Puesto 
“Pugin,” Algarrobito, Mendoza Prov., 620 
m, 4 6, 7 @, Ex: M. australis, 24-31 Jul 
1959; San Rafael, Mendoza Prov., 86, 6 @, 
Mar—May 1939; Santa Rosa, Mendoza 
Prov., 66, 7 2 (paratypes), Ex: M. aus- 
tralis, Jul & Feb 1939; Fortin Uno, Rio Ne- 
gro Prov., 1 6 (holotype), 1 2 (paratype), 
Ex: M. australis, Aug 1937; near General 
Roca, Rio Negro Valley, Rio Negro Prov., 
1 36,2 2, Ex: Ctenomys sp. Blainville, 25 
Mar 1952 (BMNH). Nacunan, Mendoza 
Prov., 1 2, Ex: Akodon sp. Meyen, 15 Mar 
1983 (REL). General Acha, La Pampa 
Prov., 1 36, Ex: M. a. australis, 20 May 
1936; La Cristina, 1 2, Ex: Galea sp. Mey- 
en, 26 Oct 1966 (USNM). 

Remarks.—All records of this species are 
restricted to Argentina, the preferred host 
being mountain cavies (M. australis). Al\- 
though this flea has not been associated 
with H. cypha, it parasitizes the same host 
species in the same general geographic re- 
gion. 


Hectopsylla gracilis Mahnert 1982 
(Figs. 10, 21) 


Material examined.—ARGENTINA: 
Puesto “‘La Carpa,’’ Mendoza Prov., elev. 
600 m, 1 2, Ex: Canis familiaris Linn., 17 
Jul 1959; Puesto “‘La Carpa,’’ Mendoza 
Prov., elev. 600 m, 6 2, Ex: G. griseoflavus 
ssp., 14 Jul 1959 (BMNH). Puerto Madryn, 
Chubut Prov., 1 d (holotype), Ex: Elig- 
modontia morgani Allen, 23 Apr 1978; 
Puerto Madryn, Chubut Prov., 1 6, 1 
(paratypes), Ex: E. morgani, 5 Apr 1978 
(Md’HN); Puerto Madryn, Chubut Prov., 1 
2, Ex: E. morgani, 24 Apr 1978 (REL). 

Remarks.—Specimens from Mendoza 
Province (BMNH) were not recognized as 
a H. gracilis until Mahnert’s description in 
1982. The geographic range of the preferred 
host, E. morgani, is limited to Argentina 
and adjacent southern Chile (Wilson and 
Reeder 1993). 


Hectopsylla knighti Traub and Gammons 
1950 
(Fig. 11) 


Material examined.—-México Michoa- 
can State, Municipality of Tancitaro, Tan- 
citami, 1 2 (holotype), Ex: “‘head of 
swift,” May 1940 (FMNBH). 

Remarks.—Additional observations of 
‘‘swifts’’ in the region of Michoacan, Méx- 
ico, are needed to find the males of this 
species and further elucidate its taxonomic 
status. 


Hectopsylla pascuali Beaucournu and 
Alcover 1990 
(Figs. 9, 20) 


Material examined.—ARGENTINA: Rio 
Chapelco, Neuquén Province, 1d (holo- 
type), 1 @ (allotype), 1 2 (paratype), Ex: 
Chelemys macronyx (Thomas), 9 Dec 1987 
(JCB). 


Hectopsylla psittaci Frauenfeld 1860 
(Fig. 22) 


Material examined.—ARGENTINA: 1 2 
(lectotype), 2 2 (paralectotypes), Ex: Strix 
pelate = Tyto alba (Scopoli); Buenos Aires, 
2 2, Ex: “owl,” 1913; Chivilcoy, Buenos 
Aires Province, 2 9, Ex: “‘hen,’’ 2 Feb 
1936 (BMNH); Las Roses, (B.A.), 1 2, Ex: 
‘“‘Paloma”’; Zapallar, La Rioja Prov., 3 °, 
Oct 1933 (USNM). BRAZIL: Progne, Rio 
de Janeiro State, 2 2 (USNM). CHILE: St. 
Jogo (Santiago de Chile), 2 2 (syntypes), 
Ex: Cyarzolyscus patagonius or Enicogna- 
thus leptorhynchus (King) (BMNH); San- 
tiago, 1 2, 11 Sep 1951; Santiago, 2 2, Ex: 
“turtle dove’? (USNM). HOLLAND: den 
Haag, 22 ¢, Ex: Phasianus sp. (L.), Aug 
1926; Rotterdam, 4 2, Ex: Gallus domes- 
ticus = Gallus gallus (L.), July 1936 
(BMNH); den Haag, 3 2, Ex: Phasianus 
sp., Aug 1926 (USNM). ENGLAND: Lon- 
don (Zoological Gardens), 3 9, Ex: Citto- 
cincla macrura (sic) = Copsychus mala- 
baricus (Swinhoe); London (Zoological 
Gardens), 3 6, Ex: ‘“‘birds in western avi- 
ary’; London (Zoological Gardens) 3 °, 


2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Ex: Cittocincla macrura (sic) or Copsychus 
saularis (L.) (BMNH); London (Zoological 
Gardens), 2 2, Ex: ‘‘Cittocincla (sic) = Kit- 
tocincla (Gould) or Copsychus’”> (USNM). 
PERU: Ex Ribeyro;sLimas lsd. 2697 Ex 
‘“‘“chickens’’; Hacienda Las Vegas, 30 km 
from Barranca City, 1 2, Ex: swallow “‘va- 
riety called Santa Rosita,’ 1947 (BMNH); 
Cuzcon, Ocongata River, elev. 3,500—4,000 
m, 2 2, Ex: Merganetta leucogenis turneri 
= Merganetta armata Gould, 26 Jul 1949; 
Hacienda Ceapano, Ocongate, Cuzco Prov., 
1 2, Ex: Colaptes rupicola puna = Colap- 
tes rupicola d’Orbigney, 14 Aug 1950 
(USNM). UNITED STATES: Oceanside, 
San Diego County, California, 1 ?, Ex: 
Bubo virginianus pacificus = Bubo virgi- 
nianus (Gmelin), 7 Jun 1942 (BMNH); 
Oceanside, San Diego County, California, 1 
2, Ex: B. v. pacificus, 7 Apr 1942 (USNM); 
Alameda (6.5 km w. Newark), California, 1 
36, 4 2, Ex: cliff swallow nests, 15 Jan 
1981 (REL). 

Remarks.—Hectopsylla psittaci, a pri- 
mary parasite of birds, is recorded from di- 
verse areas. However, it is native to the 
southern cone of South America with pop- 
ulations dispersed to the southwestern Unit- 
ed States by owls and other migratory bird 
species. Schwan et al. (1983) reported large 
numbers of this species from the nests of 
Cliff Swallows (Hirundo pyrrhonota Vieil- 
lot) and Black Phoebes [Sayornis nigricans 
(Swainson)] in southern California. The 
species likely occurs through Central Amer- 
ica as well. 


Hectopsylla stomis Jordan 1925 
(Figs. 3, 13) 


Material examined.—ARGENTINA: 
Mariano, Buenos Aires Prov., 1 2 (holo- 
type), Ex: “‘bird’’, Dec 1912; Canada Mar- 
iano, Buenos Aires Prov., 4 2 (paratypes), 
Ex: ‘‘birds,’’ 30 Dec 1912; Bahia Blanca, 
Euenos Aires Prov., 19 2 (paratypes), Ex: 
Mephitis sp. E. Geoffroy and G. Cuvier 
(sic), only Conepatus spp. Gray occur in 
Argentina, 15 Jan 1911; Pique, Buenos Ai- 
res, 1 3, 4 2, Ex: Lagostomus maximus 


VOLUME 102, NUMBER 3 


(Desmarest), 5 May 1928; El Quebrachal, 
Salta: Proy., elev: 620 m; 2 9, Ex: L. mmax- 
imus, 30 Jul 1940; El Ojito, Santiago del 
Estero Prov., elev. 620 m, 1 2, Ex: L. max- 
imus, 19 Jul 1940; La Paz, Mendoza Prov., 
elev, 620'm:"6" 6, 22 2) Ex) maximus, 1 
Mar 1939; Puesta “‘pugin’”’ Algarrobito, 
Mendoza Prov., elev. 620 m, 4 ¢, Ex: L. 
maximus, 25 Jul 1959; San Rafael, Men- 
doza Prov., 1 2, Ex: M. australis, 20 Apr 
1939 (BMNH). Chasic6, Buenos Aires 
Prov., 1 2, Ex: L. m. maximus, May 1968; 
Patagonia, 4 2, Ex: ‘‘Viscacha’’ Lagosto- 
mus sp., Sep 1962; Victoria, La Pampa 
Broviil): SsoBxeeRattus sp.y29 Jum-1935 
(USNM). 

Remarks.—Lagostomus maximus, occur- 
ring in southern Paraguay, and in northern 
and central Argentina is the primary host 
for this flea. Hectopsylla stomis has been 
collected throughout the year. 


Hectopsylla suarezi C. Fox 1929 
(Fig. 16) 


Material examined.—ECUADOR: Gua- 
moti, Chimborazo Prov., 1 6, Ex: “‘guinea 
pig’; Guamoti, Chimborazo Prov., 1 2, Ex: 
‘“suinea pig,” 8 Aug 1927. PERU: near 
Hauncabamba, Dept. Piura, 4 6, 4 @, Ex: 
Cavia aperea Erxleben (domesticated), 
1946/47; near Hauncabamba, Dept. Piura, 1 
°, Ex: ‘‘among human clothes’? (BMNH). 
Near Hauncabamba, Dept. Piura, 1 @ 
(USNM). 

Remarks.—This flea is associated with 
domestic guinea pigs associated with hu- 
man habitations. Astute collecting may 
yield many more specimens than are cur- 
rently known, although the wide use of 
DDT in human dwellings for controlling 
the vectors of Chagas’ disease (conenose 
bugs) and malaria (mosquitoes) may ex- 
plain the absence of these fleas in collec- 
tions after the mid-1940s. 


ACKNOWLEDGMENTS 


The authors express their appreciation to 
Theresa Howard, The Natural History Mu- 
seum, London; Nancy Adams, National 


623 


Museum of Natural History, Washington, 
D.C.; Robert E. Lewis, Iowa State Univer- 
sity, Ames; Jean-Claude Beaucournu, La- 
boratoire de Parasitologie Medicale, 
Rennes, France; and personnel from the 
Field Museum of Natural History, Chicago, 
and the Muséum d’Histoire Naturelle, Bel- 
gium, for the loan of specimens. 


LITERATURE CITED 


Anduze, P. J., Vogelsang, E. G., and C. F Pifano. 1947. 
Nomina de artr6podos vulnerantes actualmente 
conocidos en Venezuela. Boletin de Entomologia 
Venezolana, (nim. extraord.): 1—10. 

Augustson, G. FE and L. C. Ryan. 1948. The flea genus 
Rhynchopsyllus in the United States (Siphonap- 
tera). Bulletin, Southern California Academy of 
Science 47(3): 111-112. 

Baker, C. F 1904. A revision of American Siphonap- 
tera, or fleas, together with a complete list and 
bibliography of the group. Proceedings of the 
United Staes National Museum 27(1361): 365— 
469. 

Barrera, A. and C. Diaz-Ungria. 1957. Sifonapteros de 
Venezuela, consideraciones generales, catalogo 
anotado y claves, con exposicion grafica de los 
caracteres en ellas utilizados (Insecta: Siphonap- 
tera). Memoria de la Sociedad de Ciencias natur- 
ales la Salle, pp. 160-173. 

Cova Garcia, P. and E. Tallaferro. 1959. Pulgas mas 
comunes de Venezuela. Archivos Venezolanos de 
Patologia Tropical y Parasitologia Medica 3: 327— 
348. 

Cunha, A. 1914. Contribuigao para o estudo dos Si- 
phonapteros do Brasil, Rio de Janeiro, Rodrigues 
et Cia. 212 pp. 

Dalla Torre, C. G. 1924. Aphaniptera. Sonderabdruck 
aus den berichten des naturwisssenschaftlich-med- 
izinischen Vereines in Innsbruck 39: 1—28. 

Deignan, H. G., R. A. Paynter, Jr., and S. D. Ripley. 
1964. Check-list of birds of the world, a contin- 
uation of the work of James L. Peters, Vol. X. Jn 
Mayr, E. and R. A. Paynter, Jr., eds., Museum of 
Camparative Zoology. Cambridge, Massachusetts, 
502 pp. 

Ewing, H. E. 1929. A Manual of External Parasites. 
Charles C. Thomas, Publisher, Baltimore, Mary- 
land, 225 pp. 

Frauenfeld, G. R. von. 1860. Hectopsylla psittaci n. 
gen., n. sp. Sitzungsberichte der Akademie der 
Wissenschaften, Mathematische-Naturwissen- 
schaftliche Classe Wein 40: 462—465. 

Fuller, H. S. 1942. Notes on Neotropical Siphonaptera. 
Revista Entomologica 13(1—2): 39—44. 

Guimaraes, L. R. 1940. Notas sobre Siphonaptera e 
redescricao de Poligenis occidentalis (Almeida 


624 


Cunha, 1914). Archivos de Zoologia do Estado de 
Sao Paulo 2: 216—50. 

Haller, G. 1880. Rhynchopsyllus, eine neu Puliciden- 
Gattung, in einigen Worten gekennzeichnet. Ar- 
chiv fiir Naturgeschichte 46(1): 72—87. 

Hopkins, G. H. E. and M. Rothschild. 1953. An illus- 
trated catalogue of the Rothschild collection of 
fleas (Siphonaptera) in the British Museum (Nat. 
Hist.), Vol. I. Tungidae and Pulicidae. British Mu- 
seum (Natural History), 361 pp. 

Johnson, P. T. 1957. A classification of the Siphonap- 
tera of South America with descriptions of new 
species. Memoirs of the Entomological Society of 
Washington, No. 5, 298 pp. 

Jordan, K. 1934. On some Siphonaptera from Argen- 
tina. Revista Sociedad Entomologica Argentina 
6(1): 19-21. 

. 1939. On five new Siphonaptera from the Re- 
public of Argentina. Novitates Zoologicae 41: 
292-303. 

Jordan, K. and N. C. Rothschild. 1906. A revision of 
the Sarcopsyllidae, a family of Siphonaptera. 
Thompson, Yates and Johnston Laboratories Re- 
port (new series) 7: 15—72. 

Macchiavello, A. 1948. Siphonaptera de la costa sur- 
occidental de America (primera lista y distribu- 
cion zoo-geografica). Oficina Sanitaria Panameri- 
cana Publication 237, pp. 1—49. 

Méndez, E. 1977. Mammalian-Siphonapteran associa- 
tions, the environment and biogeography of mam- 
mals of southwestern Colombia. Quaestiones En- 
tomologicae, 13(2): 91-182. 

Peters, J. L. 1934. Check-list of Birds of the World, 
Vol. Il, Harvard University Press, Cambridge, 
Massachusetts, 401 pp. 

. 1940. Check-list of Birds of the World, Vol. 

IV, Harvard University Press, Cambridge, Mas- 

sachusetts, 291 pp. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Pinto, C. 1930. Arthrépodes parasitos e transmissores 
de doengas. Chapter 10, Siphonapteros, pp. 281— 
395. In Tratado de Parasitologia, 4, Pimento de 
Mello and Cia, Rio de Janeiro, 845 pp. 

Schreiter, R. and R. C. Shannon. 1927. Un nuevo e 
interesante género y especie de una pulga del mur- 
ciélago. Boletin Museo de Historia Natural, Univ- 
ersidad de Tucuman 1(12): 1—15. 

Schwan, T. G., M. L. Higgins, and B. C. Nelson. 1983. 
Hectopsylla psittaci, a South American sticktight 
flea (Siphonaptera: Pulicidae), established in cliff 
swallow nests in California, USA. Journal of 
Medical Entomology 20(6): 690—92. 

Sibley, C. G. and B. L. Monroe, Jr. 1990. Distribution 
and taxonomy of birds of the world. Yale Univer- 
sity Press, New Haven and London, 1,111 pp. 

Tamsitt, J. R. and I. Fox. 1970. Records of bat ecto- 
parasites from the Caribbean region (Siphonap- 
tera, Acarina, Diptera). Canadian Journal of Zo- 
ology 48(5): 1093-1097. 

Taschenberg, O. 1880. Die Fl6hé. Die arten derinsec- 
tenordnung Suctoria nach ihrem chitinskelet mon- 
ographisch dargestellt, 122 pp. 

Tipton, V. J. and C. E. Machadoo-Allison. 1972. Fleas 
of Venezuela. Brigham Young University Science 
Bulletin (Biological Series) 42(6): 1-115. 

Tipton, V. J. and E. Méndez. 1966. The fleas (Siphon- 
aptera) of Panama, pp. 289-385, plates 47—93. In 
Wenzel, R. L. and V. J. Tipton, eds., Ectoparasites 
of Panama, Field Museum of Natural History, 
Chicago, 861 pp. 

Traub, R. and J. G. Gammons. 1950. Two new fleas 
of the family Tungidae. Journal of Parasitology 
36(3): 1-4. 

Wilson, D. E. and D. M. Reeder. 1993. Mammal Spe- 
cies of the World, a Taxonomic and Geographic 
Reference, 2"! ed. Smithsonian Institution Press, 
Washington, 1,206 pp. 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 625-633 


THE PUPAE OF THREE SPECIES IN MINOMYIA THEOBALD FROM NEPAL 
AND A KEY TO THE KNOWN PUPAE OF THE GENUS IN THE ORIENTAL 
AND AUSTRALASIAN REGIONS 
(DIPTERA: CULICIDAE) 


RICHARD E DARSIE, JR. 


Florida Medical Entomology Laboratory, University of Florida, 200 9" Street, SE, Vero 
Beach, FL 32962, U.S.A. (e-mail: rfd@ gnv.ifas.ufl.edu) 


Abstract.—Four species of the genus Mimomyia are known from Nepal, Mi. chamber- 
laini, Mi. luzonensis, Mi. hybrida and Mi. intermedia. Complete descriptions and illustra- 
tions of the first three species are provided. A key to the known pupae of Mimomyia in 
the Oriental and Australasian regions is included. 


Key Words: Mimomyia, pupae, Nepal 


Four species of the genus Mimomyia 
Theobald are known from Nepal. The first 
to be reported was Mi. (Etorleptiomyia) lu- 
zonensis (Ludlow) by Pradhan and Darsie 
(1989); then Mi. (Mimomyia) chamberlaini 
Ludlow and Mi. (Mim.) hybrida (Leicester) 
by Darsie and Pradhan (1990); and most re- 
cently Mi. (Mim.) intermedia Barraud, a 
single female in the Nepal collection by 
Darsie et al. (1992). Immature stages of the 
latter are unknown. One of the purposes of 
this study is to give a complete description 
of the pupae of the former three species. 

Due to the size of the trumpets and forms 
of the paddles of the three pupae being de- 
scribed, only these structures have been il- 
lustrated and/or briefly described by Bar- 
raud (1934), Mattingly (1957, 1971), Del- 
finado (1966, Mi. chamberlaini and Mi. lu- 
zonensis)) and Baisas (1974, Mi. luzonensis 
only). Chen and Lien (1956) described the 
pupae of Mi. luzonensis and Mi. chamber- 
laini, as Mi. metallica (Leicester) in some- 
what more detail; however, no complete de- 
scription and illustration has previously 
been made. Mimomyia chamberlaini and 
Mi. metallica were considered by Mattingly 


(1957) as the same polymorphic species, 
and Knight and Stone (1977) listed Mi. me- 
tallica as an infrasubspecies of Mi. cham- 
berlaini. However, Lee et al. (1988) have 
raised Mi. metallica to a subspecies of Mi. 
chamberlaini. 

The genus Mimomyia consists presently 
of three subgenera and 45 species. Of those, 
32 are in Subsaharan Africa, mostly in the 
subgenera Mimomyia Theobald and Ingra- 
mia Edwards, while 14 occur in the Ori- 
ental and Australasian regions, 7 in the sub- 
genus Mimomyia Theobald, 4 in the sub- 
genus Etorleptiomyia Theobald, and 3 in 
the subgenus /ngramia (Knight and Stone 
1977, Knight 1978, White 1974). Pupae of 
10 species, including the three treated here, 
have been adequately described and are in- 
cluded in the following key (Belkin 1962; 
Knight and Chamberlain 1948; Mattingly 
HOSA ao). 


METHODS AND MATERIALS 


The methods are as described by Darsie 
(1998). Morphological nomenclature fol- 
lows Harbach and Knight (1980). The pu- 
pae of Mi. chamberlaini, collected in Nepal, 


626 


are in the author’s collection at the Florida 
Medical Entomology Laboratory, Vero 
Beach, FL, USA. For the other two species, 
no pupae from Nepal are available and 
those from Thailand, Malaysia, and the 
Philippines were borrowed from the Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, DC. Abbre- 
viations used in the descriptions are br 
meaning branches and Le and Pe meaning 
larval and pupal exuviae. 


DESCRIPTIONS 


Mimomyia (Mimomyia) chamberlaini 
Ludlow 
(Fig. 1) 


Pupa.—Position and size of setae as fig- 
ured, range and modal number of branches 
in Table 1. Abdomen 3.01—4.21, * 3.86 
mm, paddle 1.07—1.26, x 1.14 mm. 

Cephalothorax: Striped and mottled with 
darker brown pattern, those on mesothorac- 
ic wing match adult wing venation. Setae 
1,3,5-CT very long and stout, usually dou- 
ble; 8-CT long, mostly with 5 br (4—6); 
trumpet very long, 1.53—2.09, x 1.96 mm, 
index 12.7—17.0, * 15.6, pinna in apical 
0.16—0.23 * 19.0, with normal opening for 
water surface respiration, tracheoid part 
0.72-0.75 x 0.73 of total length. 

Abdomen: Segments I-III medium 
brown, IV—VIII with sublateral dark brown 
stripes. Seta 1-I subdendritic float seta with 
13—38 br; 2,3-I long, single; 1-II long, rath- 
er stout, with 5-10 br; 1-III-VII and 5-IV- 
VII stout, with short and long br, 1-III,1V 
with 6—12 br, 1-V,VI with 4—11 br, 1-VI 
with 4—6 br, 3—VII absent; 5-IV-VI with 4— 
11 br, 5-VII with 3—7 br; 6-III-VI long, sin- 
gle, seldom double; 6-VII ventral, short, 
usually double (1—4); 9-VIII attached to a 
rather pointed, apicolateral process, short, 
usually 3, 4 br (1—4); female genital lobes 
with rather pointed prolongation apicolater- 
ally, with medium-sized spicules. 

Paddle: uniformly brown, except for large 
white area laterally in basal 0.58—0.74, some 
pupae (70%) also with smaller white area 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


medially; index 1.75—2.18, x 1.99; midrib 
extending to apex; outer margin with large 
spine-like spicules in apical 0.2—0.44, tiny 
spicules on apex in Nepal specimens, bare 
point apically in Thai specimens; inner mar- 
gin with similar large spicules in apical 
0.12—0.22; seta 1-P short, single. 

Material examined.—NEPAL: Lumbini 
District, Shivapur, IX-13-92, 1 2 LePe, IX- 
16-92, 1 2 LePe, ex Eichornia pond (Dar- 
sie and Courtney). MALAYSIA: Perak, 
Tanjong Tualang ER., 1968, 3 2 Pe, 2 d 
Pe (811-15, 102, 104, 105, 113). PHILIP- 
PINES: Mindoro Island, San Jose, I[-25-45, 
1 3 LePe, ex Pistia marsh (E.S. Ross). 


Mimomyia (Etorleptiomyia) luzonensis 
Ludlow 
(Fig. 2) 

Pupa.—Position and size of setae as fig- 
ured, range and modal number of branches 
in Table 2. Abdomen 3.61—3.87, x 3.72 mm, 
paddle 1.0—1.04, x 1.03 mm. 

Cephalothorax: Uniformly light brown, 
setae 1,3-CT long, single, 3-CT stout; 5-CT 
medium long, usually double; 6-CT light in 
color, very long and stout, single; 8-CT un- 
usually stout, single; trumpet extremely 
long, length 2.78—3.28, x 3.03 mm; index 
25.9-35.4 xX 32.4; tracheoid 2.14—2.63, x 
2.36 and pinna 0.13—0.16 x 0.148 of total 
length. 

Abdomen: Uniformly light tan; Seta 1-I 
medium long, single; 2,3-I long to very 
long, rather stout, single; 1-II long, stout, 
double, sometimes single; 1-III-VI long, 
stout, aciculate, usually 4 br, 1-VII very 
long, triple; 3-VII absent; 5-IV-VI long, 
stout, aciculate, usually triple, 5-VII 4,5 br; 
6-II-VI long, single; 6-VII long, 4—6 br; 9- 
II-VII short, larger than usual, single; 9- 
VIII medium long or long, aciculate, 7—11 
br; female genital lobe produced apicolater- 
ally, with many large spines. 

Paddle: light tan, linear, index 4.68—5.35 
x 4.98, midrib extending to apex, outer mar- 
gin with spicules coarse in apical 0.5, finer 
basally; inner margin with coarse spicules 
to near base, seta 1-P short, thin, single. 


VOLUME 102, NUMBER 3 627 


Fig. 1. Pupa of Mimomyia (Mim.) chamberlaini. A, Cephalothorax. B, Metanotum and abdomen. C, Lateral 
process of female genital lobe. D, Enlarged spicules on border of paddle, upper from Nepal, lower from Thailand. 
Abbreviations: Bu = external buttress; CT = cephalothorax, GL = genital lobe, Mr = midrib. Pa = paddle, T 
= respiratory trumpet; scale in mm. 


628 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 1. Setal branching of Mimomyia chamberlaini pupa. 
Abdominal Segments 
Cephalo- 

Seta thorax I I Ul IV Vv VI Vil Vill 
0) — — 1 | l 1 l 

| 24 (2)! 13-38(?) 6-10(6) 7-12(10) 6-11(8) 411(7) 48 (8) 4-6 (?) — 

2 1-3 (2) 1 1 ] 1 1 1 — 

3 24(2) 1 ILC) 1-5 (2) 2-6 (3) 1-3 (2) 1-4 (3) — — 

4 1-5 (3) 3-5 (4) 2-6 (3) 1 1-5 (1) 1-3 (1) 1-2 (1) 1-4 (1) 
5) 1-5 (4) 1-3 (1) 2-8 (4) 3-6 (4) 4-10 (?) 5-11 (9) 4-10(?) 3-7 (4) — 

Gj Il 1-4 (2) 14 (1) 1 1 1251) 2 C1) 1-4 (2) — 

7 1 I) (G))) 1EORCL) l PG) 1e27 1) E22 (GL) als) — 

8 4-6 (5) 2 (Cy) = 1-3 (1) 1-3 (1) 1-3 (1) 1-3 (1) 1-4 (3) — 

9 14(@) — ] 1 1 1 1 1 14 (3) 
10 2-4 (4) — — 1 2 (CL) 12) 1 1-3 (1) _— 
11 1 (1) == = 1 1 1 1 1,2 (1) — 
WA NY) = = = = = — 1 — 
14 -— _— = 1 1 1 1 1 1 


' Range followed in parenthesis by the mode. 


Material examined.—THAILAND, Chon- 
buri Province, Bang Lamung, Khao Mai 
Keao, X-8-63, 2 2, 3 6, Pe, ex pond on 
trackway (Kol, Vuth, Chmnong; Nepal: 
Sunsari District, Tarahara XI-25-87, 1 &, 
SPP, RFD; Sindhuli Garhi District, Bardi- 
agoth, IV-28-90, 2 2, SPP. 


Mimomyia (Mimomyia) hybrida 
(Leicester) 
(Fig. 3) 


Pupa.—Position and size of setae as fig- 
ured, range and modal number of branches 
in Table 3. Abdomen 2.92-—3.47, x 3.27 mm, 
paddle 0.78—0.87, « 0.83 mm. 

Cephalothorax: Mottled with darker 
brown pattern, seta 1,3-CT very long, usu- 
ally triple; 5-CT very long, 3-5 br; 6-CT 
very long, rather stout, single; 8-CT very 
long, with 4-6 br; trumpet length 1.46—1.92, 
x 1.64 mm, index 9.0—-13.2, * 11.1; trach- 
eoid 0.82—0.1.05, * 0.93 and pinna 0.16— 
0.24 x 0.20 of total length, pinna split into 
two spinulose processes, adapted for pierc- 
ing plant tissue. 

Abdomen: Uniformly light tan; seta 1-I 
float seta, 11—16 br; stout; 1-II long, rather 
stout, with 4—7 br; seta 1-ITI-VI with thin, 
long or very long br, 1-[I with 4—9 br, 1- 
IV-VI with 2—5 br, 1-VII long, single, sel- 


dom double; 2,3-I long, single, 2-I stout; 3- 
VII absent; 5-IV-VI with thin, moderately 
long or long br, 5-I[V with 3-6 br, 5-V with 
4 or 5 br, 5-VI with 2-4 br; 6-I-VI medium 
long, single, 6-VI rarely double; 6-VII ven- 
tral, short, single or double; 9-VIII short, 
usually double, when single, aciculate; fe- 
male genital lobe with rounded projections 
apicolaterally, with numerous small spicules. 

Paddle: ovoid, entirely light brown, bi- 
lobate apically, median lobe larger, index 
1.64—-1.92, * 1.78 outer margin with coarse 
spicules in apical 0.20—0.29, inner margin 
with coarse spicules in apical 0.20—0.29, 
seta 1-P absent. 

Material examined.—THAILAND, Chieng- 
mai Province, Chiengmai, IX-30-52, 1 @, 1 
3d, LePe, ex pond (M300-A, M-300-1) (M. 
Rattanopradith); Nonthburi Province, Pak 
Kert, Ko Kret, IV-17-64, 1 92, LePe, ex 
ditch (Prajim). Nepal: Sunsari District, Tar- 
ahara, IX-X-85, 2 92; Morang District, 
Khanar, [X-X-85, 2 2; Sonapur, IX-X-85 2 
? (Burgess, unpublished data). 


KEY TO THE KNOWN PUPAE OF THE GENUS 
MIMOMYIA IN THE ORIENTAL AND 
AUSTRALASIAN REGIONS 


Partially adapted from Mattingly (1957) 


VOLUME 102, NUMBER 3 629 


at 


ee 
————e 


seed FEE 


i 


Fig. 2. Pupa of Mimomyia (Eto.) luzonensis. A, Cephalothorax. B, Metanotum and abdomen. For explana- 
tions of abbreviations, see Fig. 1. 


Ike Seta 9-VIII very large with about 8 or more with numerous branches; trumpet < 20 
br, paddle at least 4X greatest width; seta 1- width at middle (subg. Mimomyia) ...... 7 
I small, single or bifid; trumpet length at 2(1). Paddle length at least 10X greatest width; 
least 20X width at middle ............ 2, seta 1,5-VII single or bifid; trumpet with 

= Seta 9-VIII small, with at most 3 br; paddle tracheoid spiculose (subg. Ingramia) .... 3 


less than 3X greatest width; seta 1-I large — Paddle length at most 8X greatest width; 


630 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 2. Setal branching of Mimomyia luzonensis pupa. 
Abdominal Segments 
Cco)h2l> een ee 

Seta thorax I Ul i IV Vv VI Vil Vill 

0) — oo 1 l 1 ] ! 

| 1 l e252) 3,4 (4) 3,4 (4) 4,5 (4) 4-8 (5) 3 — 

2 1 l 1 ] 1 l os 

3 l 1 2h) 1-3 (3) 1-3 (2) — a 

4 1-3 (2)! 2-6 (2) l—3 (1) 14 (1) 2,3 (2) E22) 1-3 (1) 1-3 (2) 1-4 (1) 

5) 2 ] 2) 2,3 (2) 3 3 3-5 (3) 4,5 (4) — 

6 1 2G) I l l 1 1 4-6 (4) a 

7/ os (1) 2 (UY) 1,2 (1) 1-3 (2) 1-3 (1) Si) 122) = 

8 1 _ -~ Nez) l 1,2 (1) 1,2 (1) 1-3 (2) — 

9 1-3 (2) 1 l 1 1 1 l 1 7-10 (10) 
10 2 ()) _ — NG) 1,2 (2) 2s) 1-5 (1) 1-3 (1) — 
11 1 — — 2 ()) 1 1 1 - = 
12 iL? () —_— — — — — — -- -- 
14 a — — l 1 1 ] 1 1 


' Range of branching followed in parenthesis by the mode. 


seta 1,5-VII with at least 3 br, tracheoid of 
trumpet smooth (subg. Eforleptiomyia) ... 4 

. Seta 5-II,III about 0.75 length of seta 3-II- 
Ill fusca (Leicester) 

— Seta 5-IJ,HI 0.35 or less length of seta 3- 
II deguzmanae (Mattingly) 


4(2). Inner margin of paddle with small spinules 
in basal 0.5.4) ee ase es thee ees rel 5 

— Inner margin of paddle with prominent 
SPINES a: ool coe GPM Me oe ae dep e ui « 6 


. Seta 6-II, UI, VII longer than following ter- 

gum bougainvillensis (Belkin) 
— Seta 6-II, Il, VII shorter than following ter- 

gum elegans (Taylor) 
. Setae 2,3-I subequal; seta 8-II present..... 
solomonis (Belkin) 
— Seta 2-I 2.0 length of 3-I; seta 8-II absent 
luzonensis (Ludlow) 
. Trumpet with 2 pointed processes apically 


eae eater amir ears hares, a ey Mut Mo eee ae 8 
— Trumpet without 2 pointed processes at apex 
Sebiek Maine seh. Sette ey sek eee coe ere bs Y 
8(7). Setae 1,5-IV-VI with some branches longer 
than following tergum; paddle emarginate 
apicallive. cs ocr Sees hybrida (Leicester) 
—  Setae 1,5-IV-VI shorter than following ter- 
gum; paddle rounded apically .......... 
se ace esi Sieh Stussy = gurneyi (Belkin) 
9(7). Lateral pale area in basal 0.58—0.74 of pad- 


dle, 0.5 of total length chamberlaini Ludlow 
— Pale area in distal 0.5 of paddle, 0.3 of total 
length or less aurea (Leicester) 


COMPARATIVE MORPHOLOGY OF SUBGENERA 
OF MIMOMYIA BASED ON PUPAL 
CHARACTERS 


Based on evidence found in the descrip- 
tions of the pupae of 19 species in the sub- 


genus Ingramia, five species in the subge- 
nus Etorleptiomyia, and 9 in the subgenus 
Mimomyia., the latter appears to be the 
most primitive. That is because it has seta 
8-CT a normal branched seta, seta 1-I a 
dendritic float seta, and paddle with indices 
of 1.6—2.18, similar to most other mosquito 
pupae. On the other hand, the subgenera 
Etorleptiomyia and Ingramia have seta 8- 
CT stout, almost spike-like in most species, 
seta 1-I a single seta, and the paddles linear 
in shape with indices of 4.5—20. All sub- 
genera have long tracheoid respiratory 
trumpets; however, those of Mimomyia 
have indices of varying from, 9.0—17.0, 
while those of the other two subgenera have 
indices of 25—50. Contrarily, seta 9-VIII is 
greatly reduced in the subgenus Mimomyia, 
with 2—5 branches, while it is fully devel- 
oped in the other two with 8—22 branches. 
All in all, pupal evidence supports the ev- 
olutility of Etorleptiomyia and Ingramia at 
a later time than Mimomyia. These obser- 
vations also consider the work of Grjebine 
(1986) on the Ficalbiini of Madagascar. 


ACKNOWLEDGMENTS 


The author is indebted to the National Geo- 
graphic Society for partial support of this 
study; to T. Gaffigan, Walter Reed Biosys- 


VOLUME 102, NUMBER 3 631 


Bole re 


Fig. 3. Pupa of Mimomyia (Mim.) hybrida. A, Cephalothorax. B, Metanotum and abdomen. For explanation 
of abbreviations, see Fig. 1. 


632 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 3. Setal branching of Mimomyia hybrida pupa. 
Abdominal Segments 
Cephalo- 

Seta thorax I II Il IV Vv VI Vil Vill 
0) = l 1 1 l l 

1-3 (3)! 11-16 (?) 4-7 (5) 4-9 (5) 2-5 (3) 2-5 (4) 2-4 (4) 2 (A) -- 
2 Z ] 1 1 1 1,2 (1) — 
3) 14 (3) | 152, (Lb) 1-3 (1) DS(Z) 1—3 (2) — — 
4 1 (3) 2-4 (3) 1-4 (2) 3,4 (3) 2 2 2 (2) 2 (CU) 1-3 (1) 
5 3-6 (5) 1 2-356) Le (QD) 3-6 (?) 4,5 (4) 2-4 (3) 12 (2) — 
6 1 i ] 1 1 2a) 2) — 
7 1-3 @) 1 It 127 Gls) 2, (ly) 14 (1) 1 1 
8 4-6 (5) - — 2 {) 2 (Cy) i174 (Ab) 12 (Ul) 2b) ooo 
5 1-3 (1) 1 1 1 1 1 1 1 1 25@) 
10 2,3 (2) — — 1 1 125) 1 1 — 
11 il22 (ID) = — 1 1 l 1 1 -= 
12 il (U) _ — — —- — -—- — — 
14 _ — — 1 1 1 1 1 1 


' Range followed in parenthesis by the mode. 


tematics Unit, Smithsonian Institution, for 
the loan of specimens, to B. Bower-Dennis 
for the illustrations, and to Drs. J. Rey and 
C. Lord for reviewing the manuscript. This 
is Florida Agricultural Experiment Station 
Journal Series No. RO7078. 


LITERATURE CITED 


Baisas, E E. 1974. The mosquito fauna of Subic Bay 
Naval Reservation, Republic of the Philippines. 
U.S. Navy, Headquarters First Medical Service 
Wing Technical Report 72-2, 1—170. 

Barraud, P. J. 1934. Family Culicidae. Tribes Megar- 
hinini and Culicini. The fauna of British India, 
including Ceylon and Burma. Diptera. Vol. V, 463 
Pp- 

Belkin, J. N. 1962. The mosquitoes of the South Pa- 
cific (Diptera, Culicidae). I, Il, University of Cal- 
ifornia Press, 608 pp., 412 figs. 

Chen, H. and C. Lien. 1956. Mosquitoes of genus Fi- 
calbia in Taiwan (Formosa), China. Journal of the 
Formosan Medical Association 55: 199-215. 

Darsie, R. F, Jr. 1998. Descriptions of the pupae of six 
species of Armigeres Theobald, subgenus Leices- 
teria Theobald (Diptera: Culicidae) from Nepal. 
Proceedings of the Entomological Society of 
Washington 100: 234-246. 

Darsie, R. FE, Jr. and S. P. Pradhan. 1990. The mos- 
quitoes of Nepal their identification, distribution 
and biology. Mosquito Systematics 22: 69-130. 

Darsie, R. F, Jr., S. P. Pradhan, and R. G. Vaidya. 1992. 
Notes on the mosquitoes of Nepal I. New species 
records from 1991 collections. Mosquito System- 
atics 24: 23-28. 


Delfinado, M. D. 1966. The culicine mosquitoes of the 
Philippines, tribe Culicini (Diptera, Culicidae). 
Memoirs of the American Entomological Institute 
7: 1-252. 

Grjebine, A. 1986. Insectes Diptéres Culicidae Culi- 
cinae Ficalbiini. Muséum National d’ Histoire Na- 
turelle, Paris, Faune de Madagascar No. 68, 1— 
441. 

Harbach, R. E. and K. L. Knight. 1980. Taxonomists’ 
glossary of mosquito anatomy. Marlton, Plexus 
Publishing, Inc. 413 pp. 

Knight, K. L. 1978. Supplement to a catalog of the 
mosquitoes of the world. Thomas Say Foundation, 
Supplement to Volume VI. 

Knight, K. L. and R. W. Chamberlain. 1948. A new 
nomenclature for the chaetotaxy of the mosquito 
pupa, based on a comparative study of the genera 
(Diptera: Culicidae). Proceedings of the Helmin- 
thological Society of Washington 15: 1-18. 

Knight, K. L. and A. Stone. 1977. A catalog of the 
mosquitoes of the World. Thomas Say Foundation 
6: 1-611. 

Lee, D. J.. M. M. Hicks, M. L. Debenham, M. Grif- 
fiths, J. H. Bryan, and E. N. Marks. 1988. The 
culicidae of the Australasian Region. Vol. 10, En- 
tomological Monograph 2, Canberra. 

Mattingly, P. E 1957. The culicine mosquitoes of the 
Indomalayan Area Part I. Genus Ficalbia Theo- 
bald. British Museum of Natural History, pp. 1— 
6l. 


. 1971. Contributions to the mosquito fauna of 
Southeast Asia. XII. Illustrated keys to the genera 
of mosquitoes (Diptera, Culicidae). Contributions 
of the American Entomological Institute 7(4): 1— 
84. 


VOLUME 102, NUMBER 3 


Pradhan, S. P. and R. E Darsie, Jr. 1989. New mosquito 
records for Nepal. Journal of the American Mos- 
quito Control Association 5; 21—24. 

White, G. B. 1974. Priority of Ingramia Edwards, 


633 


1912, over Ravenalites Doucet, 1957, as the name 
for a subgenus of Mimomyia Theobald, 1903 
(Diptera: Culicidae). Mosquito Systematics 6: 
239-242. 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 634-642 


A STUDY OF THE GENUS FALCOSYNTRETUS TOBIAS FROM THE NEW 
WORLD WITH FIVE NEW SPECIES AND A KEY TO KNOWN SPECIES 
(HYMENOPTERA: BRACONIDAE: EUPHORINAE) 


JENO PAPP AND ScoTT R. SHAW 


(JP) Department of Zoology, Hungarian Natural History Museum, pf. 137, H-1431, 
Budapest HUNGARY; (SRS) Scott R. Shaw, Insect Museum, % Department of Renew- 
able Resources, University of Wyoming, Laramie, WY 82071-3354, U.S.A. (e-mail: 


braconid@uwyo.edu) 


Abstract.—Five new species of Falcosyntretus Tobias from North and Central America 
are described and illustrated: F. complanatus, F. falcoi, F. fallax, F. muesebecki, and F. 
transversus. A key for the six known New World species of Falcosyntretus is provided. 


Key Words: 


The genus Falcosyntretus Tobias was 
erected based on one Asian species, F. fal- 
cifer (Tobias 1965). Although a species 
now assigned to this genus, F. venustus, 
was described by Muesebeck (1936), the 
presence of this genus in the New World 
was not recognized until that species was 
reclassified by Shaw (1985). Shaw (1985) 
also established the monophyly of Falco- 
syntretus on the basis of three synapomor- 
phies: the propodeum being mostly smooth 
and polished, the petiolate first metasomal 
tergum being smooth and polished, and the 
curved ovipositor which is as long or longer 
than the first metasomal tergum. Shaw 
(1985) defined the tribe Syntretini to in- 
clude Falcosyntretus and five other genera 
sharing several synapomorphies including 
cleft tarsal claws, forewing vein M+CU ab- 
sent, and the petiolate first metasomal seg- 
ment being fused ventrally. Tobias (1986) 
indicated that the European species Syntre- 
tus xanthocephalus Marshall should be 
transferred to Falcosyntretus. Most recent- 
ly, Papp (1992) described F. elabsus from 
Korea. 

As far as known, all members of the tribe 


Falcosyntretus, Euphorinae, parasitoids 


Syntretini have distinctive cleft tarsal claws 
(Figs. 13-16) and are koinobiont endopar- 
asitoids of adult Hymenoptera, including 
bees and ichneumonids (Shenefelt 1969, 
Shaw 1988). This peculiar life style may 
partly account for their rarity in collections, 
because few attempts are made to rear par- 
asitoids from hosts such as adult bumble 
bees or adult ichneumonids. Specimens can 
be identified as Falcosyntretus using the 
key provided by Shaw (1997). A generic 
diagnosis was provided by Shaw (1985). 
Morphological terminology follows that of 
Sharkey and Wharton (1997). Types are de- 
posited at the University of Wyoming, Lar- 
amie (UWL); the Hungarian Natural His- 
tory Museum, Budapest (HNHM); the Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 
(USNM); the Canadian National Collection, 
Ottawa (CNC); and the Zoological Muse- 
um, Lund (ZML). 

The root of the generic name, falco, 
means ‘‘hawk’’ in Latin and refers to the 
falcate (curved) ovipositor of the type spe- 
cies. However, this character can be mis- 
leading for generic recognition because it is 


VOLUME 102, NUMBER 3 


clear from examination of females of sev- 
eral species that the ovipositor is very flex- 
ible, and its appearance varies depending on 
its position at death. It may appear curved 
or straight, or somewhat shorter or longer 
depending on its position or degree or ex- 
sertion at death. A more useful character for 
generic recognition is the mostly smooth 
propodeum that lacks a carinate areola 
(Figs. 9-12). 


KEY TO FEMALES OF THE NEw WorLD 
SPECIES OF FALCOSYNTRETUS 


l. First metasomal tergite distinctly narrowest 
medially, giving the appearance of a con- 
striction near the spiracles (Fig. 1); body 
size usually smaller than 3mm; antenna 
short, with 17—23 antennomeres ........ D, 

- First metasomal tergite about as broad me- 

dially as basally, or just barely narrower, but 

not appearing contricted at middle (Figs. 2— 

4); body size usually larger than 3mm, an- 

tenna longer, with more than 23 antenno- 

meres 

Occipital carina effaced dorsally (Fig. 5); 

vein cu-a of hind wing absent; vertex of 

head with a broad dark brown to black band 

that meets compound eye margins com- 

Plete lyme we asks res eo Seortes seat marr ee Oe 

Spas Falcosyntretus transversus, new species 

- Occipital carina complete dorsally (as in 
Fig. 6); vein cu-a of hind wing present; ver- 
tex of head sometimes with dark markings 
around ocelli, but never meeting margins of 
compound eyes 
a bear's Falcosyntretus muesebecki, new species 

3(1). First metasomal tergite distinctly broader 
posteriorly, surface entirely smooth (Figs. 
2—4); propodeum entirely smooth (Figs. 9— 

10) or with scattered, extremely faint rugae 
or pitting postero-medially (Fig. 11) ..... 4 

- First metasomal tergite about evenly broad 
over its entire length, not greatly broader 
posteriorly than anteriorly or medially, sur- 
face smooth but with two small longitudi- 
nally striate areas latero-medially (Fig. 4); 
propodeum with median line distinctly ru- 
gose (Fig. 12) 
Peery os Falcosyntretus falcoi, new species 

4(3). Posterior margin of median ocellus about 
even with anterior margins of lateral ocelli 
(Fig. 7); propodeum entirely smooth..... 5 

~ Posterior margin of median ocellus distinct- 
ly anterior to anterior margins of lateral 
ocelli (Figs. 6, 8); propodeum variable, 
sometimes smooth but often with faint ir- 


2(1) 


635 


regular rugae or pitting postero-medially 
(Fig. 11) .. Falcosyntretus fallax, new species 
5(4). Mesoscutum entirely black; propodeum 
with a complete semi-circular carina bor- 
dering the junction with the metasoma (Fig. 
9) .. Falcosyntretus complanatus, new species 
— Mesoscutum yellowish orange, with some 
black markings; propodeum with a distinct 
break medially in the carina bordering the 
junction with the metasoma (Fig. 10) ...... 
ete Falcosyntretus venustus (Muesebeck) 


Falcosyntretus complanatus Papp and 
Shaw, new species 
(Figs. 2, 9) 


Description of holotype female.—Body 
length 4 mm. Head in dorsal view 1.6X 
broader than long, eye somewhat protrud- 
ing and longer than temple, temple rounded 
close behind eye. Ocelli large and forming 
a low triangle, distance between median 
and lateral ocelli shorter than greatest di- 
ameter of an ocellus, posterior border of 
median ocellus even with anterior border of 
lateral ocelli. Antenna just shorter than 
body and with 30 antennomeres, first fla- 
gellomere 1.4 as long as second flagel- 
lomere, further flagellomeres proximo-dis- 
tally 2.0—2.2X as long as broad. Mesosoma 
in lateral view 1.4X as long as high. Pter- 
ostigma 3.5X as long as wide. Hind wing 
vein cu-a present basally. Hind femur 5.3 X 
as long as broad. Hind basitarsus as long as 
tarsomeres 2—3 combined. Metasoma some- 
what longer than head and mesosoma com- 
bined. Proximal 0.5 of petiole flattened, its 
hind width 1.4 greater than basal width, 
petiole itself as long as tarsomeres 1—2 
combined. Tergites long, second tergite 
slightly wider posteriorly than long medi- 
ally, third tergite quadrate or as long as 
wide behind. Ovipositor sheath as long as 
hind tarsomeres 1—2 combined. 

Color: Body color mostly golden yellow, 
darker dorsally. Antenna, longitudinal 
streak of vertex brownish black. Mesosoma 
dorsally (mesonotum, scutellum, metano- 
tum and propodeum) and petiole black. Ter- 
gites dark brown to black. Legs mostly yel- 
low, coxae and trochanters whitish, hind fe- 
mur apically and tarsomeres dark brown. 


636 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-4. First metasomal tergites of Falcosyntretus species, dorsal view. 1, F. muesebecki, 300X. 2, F. 
complanatus, 210. 3, F. venustus, 220. 4, F. falcoi, 200X. 


VOLUME 102, NUMBER 3 


Figs. 5-8. 
carina. 6, F. fallax, dorsal view, 200.7, F. complanatus, lateral view, 220. 8, F. muesebecki, dorsal view, 200. 


Wings subhyaline, pterostigma and vena- 
tion dark brown. 

Variation, paratype females.—Body 4.0— 
4.2 mm long. Head in dorsal view 1.48— 
1.54 broader than long. Antenna with 25— 
31 antennomeres. First flagellomere 1.3— 
1.4X< as long as second flagellomere. Pter- 
ostigma 3.4—3.8X as long as wide. Hind fe- 
mur 4.1—5.0 as long as broad. Hind femur 
apically and tarsomeres dark brown to 
black. 

Material examined.—Holotype ?: COS- 
TA RICA: Puntarenas, San Vito, Estac. 
Biol. Las Alturas, 1,500 m, ex. Malaise, 
January 1992 (P. Hanson) (UWL). Para- 
types: same data as holotype, 1 2 (UWL); 
same data except December 1991, 1 & 
(UWL); same data except June 1992, 2 ¢ 
(UWL); same data except March 1992, 1 @ 
(UWL); same data except May 1992, 1 @ 
(UWL); San José, Cerro de la Muerte, 26 
km. N. San Isidro, ex. Malaise, 2,100 m, 


637 


Heads of Falcosyntretus species. 5, F. transversus, dorso-lateral view, 285, note effaced occipital 


February—May 1991 (P. Hanson), 1 
(HNHM). 

Comments.—The form of the petiolate 
first metasomal tergum is distinctive in this 
species; it is broad and flat through its basal 
half, and is barely or not at all constricted 
at the middle (Fig. 2). Falcosyntretus com- 
planatus is most similar to F. venustus 
(Muesebeck) but can be distinguished by 
the presence of a complete semi-circular ca- 
rina bordering the junction of the metasoma 
to the propodeum (Fig. 9) and by the me- 
soscutum being entirely black. 

Etymology.—The species name com- 
planatus (= flattened) refers to the dorsally 
flattened petiole. 


Falcosyntretus falcoi Papp and Shaw, 
new species 
(Figs. 4, 12-13) 


Description of holotype female.—Body 
length 4 mm. Head in dorsal view 1.6 


638 


broader than long, eye somewhat protrud- 
ing and longer than temple, temple rounded 
close behind eye. Ocelli large and forming 
a low triangle, distance between median 
and lateral ocelli shorter than greatest di- 
ameter of an ocellus, posterior border of 
median ocellus even with anterior border of 
lateral ocelli. Antenna just shorter than 
body and with 30 antennomeres, first fla- 
gellomere 1.4X as long as second flagel- 
lomere, further flagellomeres proximo-dis- 
tally 2.0—2.2 as long as broad. Mesosoma 
in lateral view 1.4 as long as high. Pter- 
ostigma 3.5X as long as wide. Hind wing 
vein cu-a present basally. Hind femur 5.3 X 
as long as broad. Hind basitarsus as long as 
tarsomeres 2—3 combined. Propodeum 
mostly smooth except median line distincty 
rugulose. Metasoma somewhat longer than 
head and mesosoma combined. broad and 
flat through its entire length, parallel-sided 
with no median constriction, and entirely 
smooth except for two small longitudinally 
striate areas latero-medially. Hind width of 
petiole barely broader than anterior width. 
Entire petiole as long as tarsomeres 1—2 
combined. Tergites long, second tergite 
slightly wider posteriorly than long medi- 
ally, third tergite quadrate or as long as 
wide behind. Ovipositor sheath as long as 
hind tarsomeres 1—2 combined. 

Color: Body color mostly golden yellow, 
darker dorsally. Antenna, longitudinal 
streak of vertex brownish black. Mesosoma 
dorsally (mesonotum, scutellum, metano- 
tum and propodeum) and petiole black. Ter- 
gites dark brown to black. Legs mostly yel- 
low, coxae and trochanters whitish, hind fe- 
mur apically and tarsomeres dark brown to 
black. Wings subhyaline, pterostigma and 
venation dark brown. 

Variation, paratype female.—Essentially 
as in holotype female. 

Material examined.—Holotype 2: MEX- 
ICO: Chiapas, San Cristobal de las Casas, 
7,000 ft., 7 June 1969, Malaise trap, (CNC). 
Paratypes: UNITED STATES: 1 @, North 
Carolina, Swain Co., Smokemont, 16 July 
1977, Malaise trap, North Carolina Depart- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ment of Agriculture (USNM). HONDU- 
RAS: 1 @, Olancho, Parque Nacional La 
Muralla, 15 km N La Union, 15.07N, 
86.45W, April 1995, R. Cave, Malaise trap 
in high elevation rain forest, (ZML). 

Comments.—The form of the petiolate 
first metasomal tergum is very distinctive 
in this species: it is broad and flat through 
its entire length, parallel-sided with no me- 
dian constriction, and entirely smooth ex- 
cept for two small longitudinally striate ar- 
eas latero-medially (Fig. 4). It is most sim- 
ilar to F. complanatus but can be distin- 
guished by the mentioned petiolar 
characters, and also differs by having a dis- 
tinctly rugulose median area on the propo- 
deum (Fig. 12). The wings of this species 
were illustrated in Marsh et al. (1987, fig. 
288). 

Etymology.—This species is named in 
honor of Guido Falco, the owner of Pen- 
guin’s restaurant in Santo Domingo de He- 
redia, Costa Rica. 


Falcosyntretus fallax Papp and Shaw, 
new species 
(Figs. 6, 11, 14) 


Description of holotype female.—Body 3 
mm long. Head in dorsal view 1.7 broader 
than long, eye somewhat shorter than tem- 
ple, temple not rounded close behind eye. 
Ocelli forming a high triangle, distance be- 
tween median and lateral ocelli as long as 
greatest diameter of an ocellus, posterior 
border of median ocellus distinctly anterad 
of anterior border of lateral ocelli. Antenna 
as long as body and with 27 antennomeres, 
first flagellomere 1.3X as long as second 
flagellomere, flagellomeres proximo-distal- 
ly 1.6—2.0X as long as broad. Mesosoma in 
lateral view 1.4X as long as high. Fore 
wing as long as body. Pterostigma 3.6X as 
long as wide. Hind wing vein cu-a present 
basally. Hind femur 4.2 as long as broad. 
Hind basitarsus as long as tarsomeres 2—4 
combined. Metasoma as long as head and 
mesosoma combined. Petiole 1.5 broader 
posteriorly than basally, petiole itself as 
long as middle tarsomeres 1—2 and half of 


VOLUME 102, NUMBER 3 


Figs. 9-12. 
230X. 11, F. fallax, 110X. 12, F. falcoi, 200. 


tarsomere 3. Tergites 2—3 transverse, sec- 
ond tergite 1.4 and third tergite 2.6 as 
wide posteriorly as long medially. Ovipos- 
itor sheath as long as hind basitarsus and 
half of second tarsomere combined. 

Color: Ground color of body mostly yel- 
low. Wide longitudinal streak of vertex 
brownish black. Mesosoma above (meson- 
otum, scutellum, metanotum and propo- 
deum) and petiole black. Tergites brown. 
Legs yellow, except coxae and trochanters 
whitish. Wings subhyaline, pterostigma 
brownish opaque. 

Variation, paratype females.—Similar to 
the holotype except body length 2.6—3.1 
mm, usually 2.0—3.0 mm, long; head in 
dorsal view 1.57—-1.67X, usually 1.58— 
1.6X, as broad as long; antenna about as 
long as body and with 25-28, usually 27— 
28, antennomeres; pterostigma 3.3—3.7X, 
usually 3.5—3.7X, as long as wide; hind fe- 
mur 4.1—5.1 X, usually 4.4—4.7X, as long as 


639 


oan 


s 


Propodea of Falcosyntretus species, dorsal views. 9, F. complanatus, 210. 10, F. venustus, 


broad medially; scutellum varying from 
yellow to brown; tergites 2—7 varying from 
brown to nearly black, pterostigma varying 
from pale brown to yellowish brown. 
Material examined.—Holotype 92: 
UNITED STATES: Wyoming, Albany 
County, Medicine Bow National Forest, 5 
miles SW Lincoln Monument, mixed co- 
nifer/aspen forest, taken with Malaise trap, 
20-26 August 1990. Paratypes: 1 2, same 
data as holotype; 5 2 with same locality 
data as holotype except collected (1 each) 
on following dates: 23-38 July 1990, 28 
July—2 August 1990, 13-20 August 1990, 
15-19 July 1991, and 29 July—5 August 
1991; 1 2, same data except meadow east 
of highway, Malaise trap, 13—20 August 
1990; 2 2, same data except mixed forest 
near sagebrush, Malaise trap, 28 August 
1990; 1 2, same data except meadow near 
mixed forest, 20—27 August 1991, swept 
and aspirated, S. R. Shaw; 3 2, same data 


640 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 13-16. Cleft tarsal claws of Falcosyntretus species. 13, F. falcoi, 1,500X. 14, F. fallax, 800. 15, F. 
transversus, 1,500X. 16, F. venustus, 1,500. 


except Happy Jack Recreation Area, mixed 
forest near sagebrush, Malaise trap, 13—20 
August 1990. CANADA: 2 2, Quebec, 
Summit King Mountain, 26 June and 4 July 
1977, M. Sanborne; 1 2°, New Brunswick, 
Kouchibouguac National Park, 9 August 
1977, S. J. Miller, code 5800B; 1 3, same 
locality except 18 July 1977, G. A. Cald- 
erwood, code 5649G. Holotype and 8 par- 
atypes from Wyoming are deposited at 
UWL, four paratypes from Wyoming at 
HNHM, and the remaining (Canadian) par- 
atypes at CNC. 

Comments.—The new species, F. fallax, 
is somewhat similar to F. venustus (Mue- 
sebeck), but can be distinguished by the fol- 
lowing characters. In F. fallax the posterior 
border of the median ocellus is set distinctly 
in front of the anterior margins of the lateral 
ocelli (Fig. 6), the mesonotum is entirely 
black, the propodeum often has some irreg- 
ular rugose sculpture posteromedially (Fig. 


11), and the junction of the propodeum and 
metasoma is bordered by a semicircular ca- 
rina. In F. venustus the median ocellus is 
barely in front of the lateral ocelli, the me- 
sonotum has at least some yellow markings 
anteriorly and is often extensively marked 
with yellow, the propodeum is entirely 
smooth posteromedially, and the carina bor- 
dering the propodeal/metasomal junction is 
broken medially to form two separate short 
carinae (Fig. 10). The wings and propo- 
deum of this species were illustrated in 
Shaw (1997, figs. 19 and 58). 

Etymology.—The species name fallax (= 
false) refers to the deceptive features of this 
species that might cause it to be confused 
with F. venustus. 


Falcosyntretus muesebecki Papp and 
Shaw, new species 
(Figs. 1; 8) 
Description of holotype female.—Body 
2.7 mm long. Head in dorsal view 1.6X 


VOLUME 102, NUMBER 3 


broader than long, eye as long as temple, 
temple rounded close behind eye. Ocelli 
small and forming a high triangle, distance 
between median and lateral ocelli equal to 
greatest diameter of lateral ocellus, hind 
margin of median ocellus distinctly before 
anterior borders of lateral pair of ocelli. An- 
tenna about as long as body and with 22 
antennomeres, first flagellomere 1.2 as 
long as second flagellomere, flagellomeres 
proximo-distally 1.8—2.2 as long as broad. 
Metasoma shorter than head and mesosoma 
combined. First metasomal tergum as long 
as fore femur. Tergites 2—3 quadrate, a bit 
shorter medially than broad behind. Ovi- 
positor sheath as long as middle basitarsus 
and second tarsomere combined. 

Color: Ground color of body yellow, ex- 
cept flagellum, ocellar triangle, scutellum, 
metanotum, propodeum, and dorsum of me- 
tasoma mostly brownish black. Pterostigma 
opaque yellowish brown. 

Variation, paratype females.—Similar to 
holotype except body 2.7—2.8 mm long. 
Eye somewhat protruding and a bit longer 
than temple. Antenna about one-quarter 
shorter than body and with 19 antennom- 
eres, first flagellomere slightly longer than 
second flagellomere. Pterostigma 2.8 as 
long as wide. Hind femur 3.8 as long as 
broad medially. Ovipositor sheath some- 
times almost as long as hind basitarsus. 
Ocellar field brown to black, sometimes lat- 
eral pair of spots on mesonotum, metano- 
tum and propodeum brownish black. Hind 
half of petiole and tergites 4—7 sometimes 
dark brown, tergites 2—3 sometimes yellow. 

Variation, paratype males.—Similar to 
female except body 2.3—3.0 mm long. Head 
in dorsal view 1.65—1.7X as broad as long. 
Antenna with 17—23 antennomeres. Pter- 
ostigma 2.75—2.8X as long as wide. Hind 
femur 3.5—3.6X as long as broad. Mesoso- 
ma and metasoma brown to brownish black, 
pronotum yellow, tergites 2+3 yellow to 
brown. First metasomal tergum gradually 
wider apically, not constricted medially as 
in female. 


Material |= examined.—Holotype 2: 


641 


UNITED STATES: Georgia, Forsyth, 5—10 
June 1971, ex. Malaise trap, EF T. Naumann 
(CNC). Paratypes: 1 2, same data as ho- 
lotype (CNC); 1 female, Michigan, Onton- 
ogan County, 18 June 1960, R. and K. 
Dreisbach; 1 36, New Mexico, Catron 
County, 8 mi. S.E. Luna, 7,500 ft, 9-14 
July51979;°Ss and JeuPeck- (EN@) 1) 2 
(HNHM), 3 36 (USNM), North Carolina, 
Alleghany County, Doughton Park, Malaise 
trap operated by North Carolina Depart- 
ment of Agriculture, 25 July 1977; 1 °, 
CANADA: Newfoundland, South Branch, 
July 1973, Malaise trap, Heinrich (CNC). 

Comments.—The new species, F. mue- 
sebecki is most similar to F. transversus 
new species, which also has a small body 
length (less than 3 mm) and strongly con- 
stricted first metasomal tergum (see Fig. 1). 
Falcosyntretus muesebecki can be distin- 
guished from F. transversus by the pres- 
ence of the occipital carina dorsally, the 
presence of hind wing vein cu-a, and less 
extensive black markings on the vertex 
(black blotch not reaching compound eyes). 

Etymology.—This new species is dedi- 
cated to the late Carl E W. Muesebeck 
(1894-1987), well-known specialist of the 
parasitoid wasps and the first reviser of the 
euphorine braconids of the Nearctic Region 
(see Muesebeck 1936). 


Falcosyntretus transversus Papp and 
Shaw, new species 
(Big2 5,15) 


Description of holotype female.—Body 
2.2 mm long. Head in dorsal view 1.6X 
broader than long, eye somewhat shorter 
than temple, temple rounded continuously, 
occipital carina effaced. Ocelli small and 
forming a high triangle, distance between 
median and lateral ocelli longer than great- 
est diameter of lateral ocellus, hind border 
of median ocellus distinctly anteriad of an- 
terior borders of lateral ocelli. Antenna 
somewhat shorter than body and with 21 
antennomeres, first flagellomere hardly lon- 
ger than second flagellomere, flagellomeres 
proximo-distally 1.8—2 as long as broad. 


642 


Mesosoma in lateral view 1.4X as long as 
high. Pterostigma 3.3X as long as wide. 
Hind wing vein cu-a absent. Hind femur 
4.4X as long as broad, hind basitarsus as 
long as tarsomeres 2-3 plus half of tarso- 
mere 4. Metasoma somewhat longer than 
mesosoma. Petiole half as long as hind tib- 
ia. Tergites 2—3 transverse, second tergite 
1.6—-1.7X and third tergite 1.8—2 as broad 
as long medially. Ovipositor sheath also 
half as long as hind tibia. 

Color: Ground color of body yellow. 
Transverse streak on vertex dark brown. 
Three spots on mesonotum, scutellum, me- 
tanotum, propodeum and tergites brownish 
yelow to brown. Legs yellow, coxae and 
trochanters whitish yellow. Pterostigma 
opaque yellow. 

Variation, partype females.—Similar to 
the holotype except body 2.1mm _ long. 
Head in dorsal view 1.5 as broad as long. 
Antennae with 20—21 antennomeres. Pter- 
ostigma 3X as long as wide. Hind femur 
4.5 as long as broad. 

Material examined.—Holotype @&: 
UNITED STATES: Wyoming, Albany 
County Medicine Bow National Forest, 2 
miles N on road No. 705, willow bog, taken 
with Malaise trap, 19—23 July 1991 (UWL). 
Paratypes: 1 2, same data as holotype ex- 
cept 9-15 July 1991 (UWL). CANADA: 1 
2, Manitoba, Riding Mountain National 
Park, Dead Ox Creek, hardwood forest, 400 
m, 28 June 1979, Mason (CNC). 

Comments.—The new species, F. trans- 
versus, is distinctive by the absence of the 
occipital carina and the transverse dark 
brown streak on upper part of head meeting 
the eyes. 

Etymology.—The species name “‘trans- 
versus” refers to the transverse brown 
streak above on the head. 


ACKNOWLEDGMENTS 


We thank Dr. David Smith (Systematic 
Entomology Laboratory, USDA, Washing- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ton, D.C.) and Dr. John Huber (Canadian 
National Collection, Ottawa) for providing 
loans of specimens. Ms. Teresa Williams 
(Western Research Institute, Laramie) as- 
sisted with environmental scanning electron 
microscopy. Ms. Norma Hosher assisted 
with word-processing. 


LITERATURE CITED 


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, No. 13, 98 pp. 

Muesebeck, C. E W. 1936. The genera of parasitic 
wasps of the braconid subfamily Euphorinae, with 
a review of the Nearctic species. United States 
Department of Agriculture Miscellaneous Publi- 
cations 241: 1—38. 

Papp, J. 1992. Braconidae (Hymenoptera) from Korea, 
XIV. Acta Zoologica Hungarica 38(1—2): 63-73. 

Sharkey, M. J. and R. A. Wharton, 1997. Morphology 
and terminology, pp. 19-37. Wharton, R. A., P. 
M. Marsh, and M. J. Sharkey, eds., Manual of the 
New World Genera of the Family Braconidae 
(Hymenoptera), Special Publication of the Inter- 
national Society of Hymenopterists, Number 1, 
Washington, D.C., 439 pp. 

Shaw, S. R. 1985. A phylogenetic study of the subfam- 
ilies Meteorinae and Euphorinae (Hymenoptera: 
Braconidae). Entomography 3: 277-370. 

. 1988. Euphorine phylogeny: the evolution of 

diversity in host utilization by parasitoid wasps 

(Hymenoptera: Braconidae). Ecological Entomol- 

ogy 13:323-335. 

. 1997. Subfamily Euphorinae, pp. 234—254. In 
Wharton, R. A., PR. M. Marsh, and M. J. Sharkey, 
eds., Manual of the New World Genera of the 
Family Braconidae (Hymenoptera), Special Pub- 
lication of the International Society of Hymen- 
opterists, Number 1, Washington, D.C., 439 pp. 

Shenefelt, R. D. 1969. Hymenopterorum Catalogus, 
Braconidae 1, Euphorinae. W. Junk, The Hague. 
176 pp. 

Tobias, V. I. 1965. Generic groupings and evolution of 
parasitic Hymenoptera of the subfamily Euphori- 
nae (Hymenoptera: Braconidae), I. Entomological 
Obozrrenie. 44: 841—865 (in Russian). [Transla- 
tion in: Entomological Review Washington 44: 
494-508. ] 

. 1986. Subfamily Euphorinae, pp. 181—250.Jn 

Medvedev, G. S., ed., Identification Keys for In- 

sects of the European Part of the U.S.S.R., Vol. 

III. Hymenoptera. Part 4. Nauka Publishing 

House, Leningrad (in Russian). 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 643-651 


PREY SELECTION AND DIURNAL ACTIVITY OF 
HOLCOCEPHALA OCULATA (F.) (DIPTERA: ASILIDAE) IN 
COSTA RICA 


Louis M. LAPIERRE 


Department of Organismic Biology, Ecology, and Evolution, University of California, 
Los Angeles, CA 90095-1606, U.S.A. (e-mail: louis @ucla.edu) 


Abstract.—Holcocephala oculata (F) is an abundant small robber fly in open areas at 
La Selva Biological Station, Costa Rica, and was studied during August 1995. The ma- 
jority (92%) of prey taken by H. oculata belong to the orders Hymenoptera, Diptera, and 
Coleoptera, with the remaining belonging, in order of decreasing frequency, to Hemiptera, 
Thysanoptera, Homoptera, Araneida, Strepsiptera, and Psocoptera. A comparison of sticky 
trap samples of flying insects and robber fly prey items reveal differences in the proportion 
of orders represented. Either H. oculata prefers particular orders of prey, or sticky traps 
are inadequate in sampling insect faunas. Cannibalism was not observed for H. oculata. 
Flies are more numerous in sunny areas at the beginning and end of the day. Flies are 
rarely observed in exposed areas on clear days when incident radiation is high. Areas 
shaded from direct incident radiation through the day show little change in fly numbers. 
Courtship and copulation, described herein, generally occur after 1200. Holcocephala 
oculata perches less than a meter off the ground, on average, with grass species tending 
to be the substrate most often chosen. No evidence for invertebrate predation on H. oculata 
was found, although several suitable predaceous arthropods co-occur with the robber fly 
and at least one, the ponerine ant Ectatomma ruidum Roger, readily attacked robber flies 
during feeding trials. 


Key Words: Asilidae, Holcocephala oculata, robber fly, prey selection, courtship, diurnal 


activity, La Selva Biological Station, Costa Rica 


Robber flies are among the larger and 
more visible members of the Diptera but are 


item externally. In addition, some members 
of this group have courtship displays that 


poorly understood when it comes to details 
of their ecology and behavior. A number of 
temperate zone Asilidae have been studied 
in detail (Lavigne and Holland 1969; Hes- 
penheide and Rubke 1977; Hespenheide 
1978, 1989; Weeks and Hespenheide 1985), 
but only a few tropical species (Fisher 
1983; Shelly 1984a, 1988; Fisher and Hes- 
penheide 1992). These flies typically cap- 
ture their prey by waiting at a perch, inter- 
cepting the prey in mid-flight, and then re- 
turning to the perch and consuming the 


usually involve characteristic aerial maneu- 
vers on the part of the male (Fisher and 
Hespenheide 1982, 1992). 

Holcocephala oculata (F.) (Dasypogoni- 
nae: Danilini; Fig. 1) is the smallest (mean 
length 5.20 mm; n = 10) of four species of 
Holcocephala that are known to occur at La 
Selva Biological Station, Costa Rica (E. 
Fisher and H. A. Hespenheide, unpub- 
lished). Adults perch on low-growing veg- 
etation in open areas. Here I describe the 
prey selection and diurnal activity of H. 


644 


Fig. 1. 
perch behavior. 


Holcocephala oculata showing typical 


oculata and test the hypothesis that net in- 
cident radiation affects robber fly densities 
more than ambient temperature or time of 
day. 


MATERIALS AND METHODS 


The data were collected during 10 sepa- 
rate days between 9-29 August 1995 at the 
Organization for Tropical Studies’ La Selva 
Biological Station which is located at the 
confluence of the Rio Puerto Viejo and Rio 
Sarapiqui, Heredia Province, Costa Rica 
(McDade et al. 1994). 

I chose two 50 m transects along weedy 
margins bordering mono- and polyculture 
plots within the “‘Huertos”’ project, an on- 
going study on the sustainability of soil fer- 
tility in reconstructed tropical ecosystems 
(J. Ewel, personal communication). One 
transect was continually exposed to the sun 
from dawn until dusk (“‘sunny”’?) whereas 
the other was shaded most of the time 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


(‘“‘shady’’). In addition to the transects, 
neighboring weedy margins within the 
Huertos project were used for additional be- 
havioral and ecological observations on H. 
oculata. The weedy margins are cut at three 
to six month intervals and maintained at a 
height of less than three meters, and include 
several species of grasses, the most com- 
mon being Paspalum conjugatum Berg., 
Digitaria sp., and Cynodon dactylon (L.) 
Pers. In addition, sedges (Cyperus spp.), 
and a diverse assemblage of plant species 
adapted to disturbed areas are common 
within the margins, as are fallen stems and 
leaves from adjacent tree plots. 

At least five additional asilid species 
(three Holcocephala, one Atractia, one 
Mallophora) also co-occur with H. oculata 
in the weedy margins, and many species of 
predaceous arthropods may be found there 
as well (personal observation). The biology 
of one other species (H. affinis (Bellardi)) 
has been studied at La Selva and is gener- 
ally similar to that of H. oculata (H. A. 
Hespenheide, unpublished). 

To determine the diversity of prey taken 
by H. oculata, | captured feeding robber 
flies in the sunny transect during the 10 
days and collected their prey items. Prey 
were collected by placing a plastic vial over 
a robber fly disturbing it so that it released 
the prey item into the vial, after which the 
robber fly was released. Preliminary cap- 
tures revealed no detrimental effects due to 
this method of removing prey from the rob- 
ber flies; released individuals remained in 
the transect and were often seen with a new 
prey item moments later. This method of 
prey removal has been used by others (see 
Hespenheide and Rubke 1977; Hespenheide 
1978, 1989; Shelley 1984a, 1988). I record- 
ed the time at which the prey item was col- 
lected in order to observe any temporal pat- 
tern in prey choice. Prey were not sampled 
from robber flies in the shady transect. 

To gather information on the diversity of 
available prey items, I placed two sticky 
traps (21.5 X 28 cm transparencies coated 
with Tanglefoot®) on poles 1 m above the 


VOLUME 102, NUMBER 3 


ground in the sunny transect during study 
days 8—10 while the robber flies were being 
sampled. Only arthropods measuring up to 
4.3 mm (the maximum prey size collected 
from H. oculata) were considered in the 
analysis of the sticky traps. I identified prey 
and sticky trap samples to order. To deter- 
mine if H. oculata exhibits a preference for 
certain prey over others or is behaving op- 
portunistically, I performed two analyses to 
compare the actual robber fly prey with the 
available prey gathered from sticky traps. 
In one analysis I compared a subsample of 
the actual prey collected during the days the 
sticky trap samples were being employed 
(days 8—10). In a second analysis I included 
the entire actual prey sample collected dur- 
ing the 10 sampling days. 

To determine the daily cycle of H. ocu- 
lata, 1 censused flies on all 10 days dividing 
each into five two-hour periods beginning 
at 0800 h and ending at 1600 h. I conducted 
a census at the beginning of each of the five 
periods. Flies in the shady transect were 
simply counted while those in the sunny 
transect were counted and assigned to one 
of four activities: Feeding, copulating, 
courting, or other. The activity “‘other”’ in- 
cluded flies that were perched but not feed- 
ing and those that were flying between 
perches. During the intervening time peri- 
ods, I kept out of the transects to minimize 
my disturbing the flies. 

Prior observations on the daily cycle of 
H. oculata indicated that individuals avoid- 
ed areas exposed to direct sunlight during 
the hottest times of the day, especially on 
clear days. I gathered data on ambient tem- 
perature and incident radiation to determine 
possible influences on the daily cycle of H. 
oculata. My hypothesis was that net radia- 
tion was more important than temperature 
or time of day in determining robber fly 
density in a given area; the null hypothesis 
was that there was no difference among the 
three independent variables in determining 
robber fly density. Net radiation was mea- 
sured with a net radiometer monitored by 
the Huertos project. Ambient temperature 


645 


was measured at a single fixed location in 
the shade between the two transects. 

To describe the courtship behavior of H. 
oculata, I surveyed the transects during the 
study periods and observed courting pairs. 
I recorded the time and noted whether the 
outcome of the courting attempt was suc- 
cessful or unsuccessful in initiating copu- 
lation. 

To determine whether or not H. oculata 
exhibits a preference for type or height of 
perch, I walked the weedy margins in and 
around the transects and recorded the 
choice of perch substrate, location on sub- 
strate, and height of perch above ground for 
all H. oculata encountered. The location 
was defined as precisely where on the sub- 
strate the fly chose to perch. Above ground 
height referred to the shortest distance be- 
tween the fly’s position on the substrate and 
the ground. 

For all analyses, data on mean values are 
evaluated with the non-parametric Wilcox- 
on sign rank test, and contingency tables 
employing the G-test or Fisher’s exact test 
are used to compare percentile data. 

The arthropod predators most often ob- 
served in the transects foraging on sub- 
strates used by H. oculata were Ectatomma 
ruidum Roger, an aggressive ponerine ant, 
and orb-weaving spiders (Araneidae). I pre- 
formed feeding trials to determine the pal- 
atability of this fly to E. ruidum with robber 
flies disabled by having one wing removed. 
Orb-weavers construct webs in the weedy 
margins at the level of H. oculata’s perch 
heights. Because prey items accumulate in 
these webs, information on the spider’s prey 
choice is readily available. I carefully re- 
moved the silk from entrapped prey to es- 
tablish whether H. oculata was among 
those preyed upon by the spiders. Other ar- 
thropod predators observed included jump- 
ing spiders (Salticidae), which have been 
observed to take robber flies (H. A. Hes- 
penheide personal communication), and ti- 
ger beetles (Cicindelidae). However, I ob- 
served so few individuals (n = 4 and n = 


646 


2, respectively) that I am unable to evaluate 
their potential as robber fly predators. 


RESULTS AND DISCUSSION 


I collected nine orders of arthropod prey 
from H. oculata (Table 1). Among these the 
Hymenoptera, Diptera, and Coleoptera to- 
gether account for 92% of the total prey 
sample. The most frequent prey at the study 
site were ant reproductives (28.5%). The 
average prey size was 1.48 mm (n = 379) 
with a range of 0.4 mm (nematocerous Dip- 
tera) to 4.3 mm (Coleoptera). Hymenoptera, 
Diptera, and Coleoptera were also found to 
dominate prey samples of Holcocephala 
fusca Bromley and H. abdominalis (Say) in 
the eastern United States (Dennis 1979, 
Scarbrough 1982). However, H. calva 
(Loew), also from the eastern United States 
and while showing a similar preference for 
Hymenoptera and Diptera, differs from oth- 
er Holcocephala studied to date by prefer- 
ing Hemiptera, Homoptera, and Psocoptera 
over Coleoptera (Scarbrough 1982). The 
predator to prey size ratio for H. oculata is 
3.51 (mean robber fly length 5.20 mm). 
This is identical to that determined for H. 
fusca (3.5, mean robber fly length 5.9 mm; 
Dennis 1979), and intermediate between H. 
abdominalis and H. calva (3.4 and 3.7, re- 
spectively, mean robber fly lengths 5.81 
mm and 7.25 mm, respectively; Scarbrough 
1982). Compared to other asilid genera, the 
predator to prey size ratio for H. oculata is 
close to that reported for the similarly sized 
robber fly Nannocyrtopogon neoculatus 
Wilcox and Martin (3.66, mean robber fly 
length 6.62 mm) in the southwestern United 
States (Hespenheide1978), but differs from 
that reported for the much larger Sticho- 
pogon trifasciatus (Say) (2.23, mean robber 
fly length 13.00 mm), also from the south- 
western United States (Weeks and Hespen- 
heide 1985). No incidence of cannibalism 
by H. oculata was observed. 

Factors that contribute to prey choice for 
particular asilid species likely include the 
relative difficulty in handling prey as well 
as the availability of prey in space and time 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


(Lavigne and Holland 1969, Hespenheide 
and Rubke 1977). For H. oculata, compar- 
isons of available prey from sticky traps 
with actual prey collected during the same 
period (days 8—10) and actual prey from the 
remaining days (days 1—7) show dissimilar 
proportions with respect to some prey or- 
ders (Fig. 2). For days 8-10, Hymenoptera, 
Diptera, and Coleoptera are the predomi- 
nant orders of actual prey of H. oculata as 
well as of sticky trap samples at 90% and 
83%, respectively. In general, there is no 
significant difference between the two sub- 
samples of actual prey. This could indicate 
that H. oculata is consistent in prey choice, 
or that the relative availability of prey 
changed little over the course of the study. 
The higher proportion of formicid Hyme- 
noptera among prey could be the result of 
robber flies preferring this group. The high- 
er proportion of non-nematocerous Diptera 
in the sticky trap samples could be due to 
robber flies not favoring this group, perhaps 
because they are generally too difficult to 
capture. These results are in contrast to 
those found by Shelly (1984a) for Atractia 
marginata Osten Sacken in Panama where 
non-nematocerous Diptera composed a 
lower proportion of prey relative to what 
was available from sticky traps, and Hy- 
menoptera were insufficiently represented 
in either sample. Alternatively, the differ- 
ences observed between robber fly prey and 
sticky trap samples may be an artifact of 
the limited ability of sticky traps to ade- 
quately sample them, and of sticky traps 
themselves being more attractive to some 
taxa than others (Shelly 1984b, 1988; Men- 
sah 1996). 

Size comparisons among and between 
actual and available prey also reveal signif- 
icant differences (Table 1). In general, there 
is no difference (P > .8; Wilcoxon rank 
sum test) between the average size prey tak- 
en by H. oculata and the average size prey 
available to it. Differences emerge, how- 
ever, when comparisons are made between 
individual prey orders. These results indi- 
cate that H. oculata prefers significantly 


VOLUME 102, NUMBER 3 647 


0.3 


[| Actual prey (days 8-10) 


at 0.25 WM Available prey (days 8-10) 
: FF] Actual prey (days 1-7) 
=) 02 
2 
jor 
0.15 7 
= 
£ 
= 
SOx a 
e a 
= b 
0.05 
b aba 
Thy Hom 


S 
~) 
n 


© 
Nn 


Proportion of H. oculata 


0.25 


0800 1000 1200 1400 1600 


Time (hours) 


Figs. 2-3. 2, Proportions of actual and potential prey of H. oculata: available prey from sticky traps (days 
8-10; closed bars), actual prey from H. oculata (days 8-10; open bars), and remaining actual prey (days 1-7; 
cross-hatched bars). Relationships among taxa that are significantly different at P < .05 (G-test) are represented 
by different letters. Prey taxa are defined as follows: Hym-F ants; Hym-NE non-ant Hymenoptera; Dip-N, 
nematocerous Diptera; Dip-NN, non-nematocerous Diptera; Col, Coleoptera; Hem, Hemiptera; Thy, Thysanop- 
tera; Hom, Homoptera. 3, Proportion of H. oculata involved in various activities at each census period. Values 
for each census period are summed over the 10 study days. CP = copulating, CT = courting, F = feeding, O 
= other (i.e., perched without prey or flying). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


648 


“sasATBUR [BONSN}S OY} Ul pasn d1OM (GS) SUONeIADP plepurys jnq ‘aTqei oy} Ul payuasoid ore sIOIa prepUur}s p 
[PAI SO’ > d UY} Je JULOYIUSIS oIe Jey} SOOUdIOFJIP SOYNUSp! Ysiojse uy 

‘Iap1io Aq (Ol —g skep) Aaid ayquyieae Jo ozIs uvoul jsulese Japi1o Aq (Q[—g8 sAep) Aoid yenjoe JO 9ZIs URDUT JO 3S} UNS YURI poUsIs UOXODSTIAA Aq UOsTIedWIOD 5 
‘[2AI] SO’ > d OU) J JULOYIUSIS oe Je] SOOUDIOJJIP SOYNUOPT Yst19}se 

UY ‘PeUuTquIOS siapio [je YM Aad yenjoe Jo 9ZIS URDU [][eIOAO jsUIeSe JopIo Aq Aald Jenjoe JO 9ZIS UROUT JO Sd} WINS YURI pousIs UOXODTIAA Aq uOstIedUIOD 
‘piajdososg ‘osg ‘eiajidisdans ‘ng ‘eprouery ‘ery ‘eiajdowoy ‘woyY ‘e1ajdoursAyy, ‘Ay ‘esrajdruay ‘way ‘etojdoapoD ‘[oD ‘viaidiq sno13d90}euIdU-uOU 

NN-diq ‘esaidiq snoro.0jyeurou ‘N-diq ‘e1ajdousw AY prorutoj-uou ‘4N-wAH ‘eioydouswiAH proruiioy ‘4-WAY ‘:SMOT[OJ se poyerAciqqe ose siopio podoryuy » 


Iv 7v0 9 tee, Il 0°00! C81 ev vo 61 sr I 0-001 6LE [RIOL 
a = 7 a a az = O08 T £0 I OSd 
Ta = Po = = LOLI =d G00 Ol 080 0) G NS 
=F rar —_ 7 —, *lEvO =d Cc 1-90 Gia S60 9'1 9 RIV 
— OV-S 0 OV 60°C VS Ol Corl = d Ce OT 0c Orc 61 is wloH 
SCCOs rd L' 1-60 90° QI L6 81 OLCE, di —8'°0 fe Oc | IG 8 AUL 
a LE Sy £0" EY I ol io OcLS =d Gm Cal i) O81 iG 8 wioH 
«8900 =d OrO0T (EM (AMI te C6 LI Ivcs8' =d ev-S 0 Ol O8'1 6LI 89 19) 
KOVVO = d EW Ne cr | 6 VC OV cr8c =d Oc S 0 0) Ol oma eV NN-diq 
eSel =d Se) LO’ (GE || Eve cv v060' = d Or v0 90° 00'| 9'0C 8L N-diq 
LéL6) = d ce 0 80° 88°0 ECG CV «C810 = d CeO 90° Ol Gie) OS AN-wWAH 
9990 =d i El © 8 8CC GG V *LOVO’ = d Giv-E I 90° OST 8c 801 J-wAY 
> uosuedWOD azis asury ras urs % ‘ON yW uosueduloD 2zIsg asury pds uray % ‘ON Aeplo 
* (urut) azig (wul) azIg 
sdeay, AyONS wor Aogig [emoy pIpjNIO “FY wor Kaig jenwoy 


“S9]0UJOOF OY) UT 
poure[dxs ore g pue vy suostieduiod ozts Jo sasAyeuy ‘sofdures oy) ul payuasaidar siapio ay} 10} pajyuasoid are ozis pue ‘ajdures Jo Juaosad ‘(‘ON) Poloa[[oo Jaquinu 
ay] UO RIV “ROTY BISOD UONRIS [BOISO][OIg BATAS eT Ie Sdey AYONS UO paydaTjoo Aoid ayquyTeae pue YInjNIO “Y Aq usaye} Add [enjor JO soNSLIa}ORIVYD = “[ FGeL 


VOLUME 102, NUMBER 3 


smaller non-nematocerous Diptera prey 
than the average size available to it, and 
significantly larger Coleoptera. H. oculata 
chooses significantly larger formicid Hy- 
menoptera than the overall average size of 
prey captured, and smaller non-formicid 
Hymenoptera and Araneida. This apparent 
size preference has also been observed in 
other robber flies (Hespenheide and Rubke 
1977, Hespenheide 1978, Shelly 1984a) 

The proportion of flies involved in the 
four activities (copulation, courtship, feed- 
ing, or other) at different times of the day 
is shown in Fig. 3. Copulation and court- 
ship tend to peak in mid- to late afternoon 
and the proportion of flies feeding generally 
increases through the day. A linear regres- 
sion shows a strong inverse relationship be- 
tween fly dens'ties and net radiation (Fig. 
4; P < .000’ r*? = .54) and a weaker in- 
verse relauunship between fly densities and 
ambient  nperature (P < .0001; r° = .34), 
althougn u.ese factors are correlated (P < 
.0OO1; r2 = .67). 

Differences in robber fly behavior be- 
tween the shady and sunny transects further 
support the hypothesis that net radiation is 
the primary influence. The cycle of average 
daily robber fly density in the sunny and 
shady transects is shown in Fig. 5. During 
the study period, robber flies in the shady 
transect exhibited no significant change in 
average hourly density throughout the day 
Whereas those in the sunny transect de- 
creased significantly at midday compared to 
dawn or dusk. By 1400 h direct sunlight 
had entered the shady transect due to the 
sun’s position at that hour, and the affect of 
this is represented in Fig. 5 as the period in 
the shady transect during which the fewest 
flies were counted. Flies in the shady tran- 
sect tended to congregate in shady patches 
when sunlight entered the area, or relocate 
out of the transect. In the sunny transect, 
the few flies that chose to remain when net 
radiation was highest were found perched 
under grass blades, positioned in such a 
way so that the blade blocked the sun’s 
rays. These observations indicate that the 


649 


H. oculata density 


0 200 400 600 


Net radiation (watts/m2) 


800 


Mean H. oculata density 


0800 


1000 1200 1400 1600 


Time (hrs.) 


Figs. 4—5S. 4, Linear relationship between H. oculata 
density and net radiation: as net radiation increases fly 
density decreases (P < .0001; r°? = .54). 5, The daily 
cycle of H. oculata in the sunny transect (open circles) 
and shady transect (closed circles). Data are shown as 
the mean density, + SE, for each of five time periods 
for 10 separate days. Different letters represent signif- 
icant differences among times within transects at P < 
.O5 (Wilcoxon sign rank test); letters a—c refer to com- 
parisons within the sunny transect, and letters d and e 
to comparisons within the shady transect. 


flies may be responding primarily to inci- 
dent radiation. Robber flies in general are 
believed to relocate to more shaded perches 
presumably to maintain a more “normal” 


650 


body temperature (Lavigne and Dennis 
1973). 

Fourteen pairs of H. oculata involved in 
courtship were observed in detail during 
this study with five resulting in the male 
joining the female in copulo. Males locate 
females by slowly cruising through the hab- 
itat close to the ground, pausing briefly to 
inspect perch sites for females. When a 
perched female is located, the male hovers 
several centimeters behind and slightly be- 
low the perched female and begins to fly 
upwards and make physical contact with 
her. Each time the male makes contact, the 
female’s wings are pushed apart; possibly 
to provide him with an exposed abdomen 
to hold on to for leverage in initiating cop- 
ulation. These collisions happen, on aver- 
age, at about seven second intervals be- 
tween which the male resumes the hovering 
position, rubbing his tarsae together prior to 
the next attempt. Once in copulo the pair 
remains attached, tail to tail with the female 
perched and the male dangling upside 
down, for an average of 11 minutes (n = 
5). The female will often relocate to another 
perch, towing the male behind her. The nine 
unsuccessful courtship attempts ended with 
the female, apparently uninterested in 
courting the male, flying far enough away 
such that he lost track of her. The copula- 
tory position I describe here for H. oculata 
is similar to that of H. fusca in North Amer- 
ica (Dennis 1979). 

Among the 80 H. oculata observed dur- 
ing the study of perch preference, most 
chose grasses (90%) over woody branch 
tips, herbs, and sedges in and around the 
study area, with a mean + SE perch height 
of 45.5 + 1.7 cm. The grass species and 
their order of perch preference included P. 
conjugatum (39%), Digitaria sp. (24%), C. 
dactylon (18%), and undetermined blades 
(19%). Among the major parts of an indi- 
vidual grass, 60% of robber flies preferred 
the tips of spikes, 28% the main part of the 
blade, 9% the tip of the blade, and 3% the 
stem of the spike. This clear preference for 
grasses, and for P. conjugatum in particular, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


may be an artifact of their relative abun- 
dance being higher in these margins, which 
appears to be the case. Alternatively, H. 
oculata exhibits a species-specific perch be- 
havior (see Fig. 1) like many asilids (Fisher 
and Hespenheide 1982, 1992), and grasses 
may present the optimal substrate for this. 

Ectatomma ruidum seized the robber flies 
immediately in all 10 of the feeding trials. 
I conclude from this that, given the oppor- 
tunity under non-experimental conditions, 
the ant would prey upon H. oculata. 

Of 62 prey items removed from 18 orb- 
weaver webs, 47 were ant reproductives, 
nine were flies, and six were beetles. Inter- 
estingly, these orders also dominated the 
prey sampled from Micrathena schreibersi 
(Perty) webs in Panama (see Shelly 1984b). 
None of the orb-weaver’s prey in my study 
were robber flies. The absence of robber 
flies as orb-weaver prey might be a result 
of robber flies cruising through the habitat 
at a slow enough rate as to evade becoming 
ensnared. I observed one case in which a 
robber fly collided with a web and imme- 
diately flew around it. Non-nematocerous 
Diptera are strong fliers and as a group have 
been found to be disproportionately repre- 
sented in webs of M. schreibersi (Shelly 
1984b), and this may likely be due to their 
ability to both avoid and escape becoming 
caught. This may explain the absence of H. 
oculata in orb-weaver webs during my 
study. 


ACKNOWLEDGMENTS 


I am grateful to Seth Bigalow, Ankila 
Hiremath, and John J. Ewel for allowing me 
to work in the study site of the Huertos pro- 
ject (NSF grant DEB-9318403) and for pro- 
viding the data on net radiation. I am also 
grateful to the principal investigators and 
staff of the Arthropods of La Selva project 
(John T. Longino, Robert K. Colwell, Dan- 
ilo Brenes Madrigal, Ronald Vargas Castro, 
Maylin Paniagua Guerrero, and Nelci 
Oconitrillo Miranda; NSF grant DEB- 
9706976) for allowing me to use the facil- 
ities contained there. I thank Grant L. Gen- 


VOLUME 102, NUMBER 3 


try for first introducing me to H. oculata, 
and Henry A. Hespenheide and A. G. Scar- 
brough for reviewing the manuscript and 
offering valuable comments. Eric Fisher 
provided the final determination for H. ocu- 
lata. Funding support during this project 
was supplied by the Vice-Chancellor for 
Research at UCLA, Kumar Patel, and the 
Organization for Tropical Studies (OTS). 


LITERATURE CITED 


Dennis, D. S. 1979. Ethology of Holcocephala fusca 
in Virginia (Diptera: Asilidae). Proceedings of the 
Entomological Society of Washington 81(3): 366— 
378. 

Fisher, E. M. 1983. Pilica formidolosa (mosca asesina, 
robber fly), pp. 755-758. In Janzen, D. H., ed., 
Costa Rican Natural History. The University of 
Chicago Press, Chicago, 816 pp. 

Fisher, E. M. and H. A. Hespenheide. 1982. Taxonomy 
and ethology of a new Central American species 
of robber fly in the genus Glaphyropyga (Diptera: 
Asilidae). Proceedings of the Entomological So- 
ciety of Washington 84(4): 716-725. 

1992. Taxonomy and biology of Central 
American robber flies with an illustrated key to 
genera (Diptera: Asilidae), pp. 611—632. /n Quin- 
tero, D. and A. Aiello, eds., Insects of Panama and 
Mesoamerica: Selected Studies. Oxford Universi- 
ty Press, New York, New York, 692 pp. 

Hespenheide, H. A. 1978. Prey, predatory and court- 
ship behavior of Nannocrytopogon neoculatus 
Wilcox and Martin (Diptera: Asilidae). Journal of 
the Kansas Entomological Society 51(3): 449— 
456. 

. 1989. Notes on the biology of Efferia wilcoxi 

(Bromley) (Diptera: Asilidae). The Pan-Pacific 

Entomologist 65(4): 375-380. 


651 


Hespenheide, H. A. and M. A. Rubke. 1977. Prey, 
predatory behavior, and the daily cycle of Holo- 
pogon wilcoxi Martin (Diptera: Asilidae). Pan-Pa- 
cific Entomologist 53(4): 277-285. 

Lavigne, R. J. and E R. Holland. 1969. Comparative 
behavior of eleven species of Wyoming robber 
flies (Diptera: Asilidae). University of Wyoming 
Agricultural Experimental Station Science Mono- 
graph 18: 1-61. 

Lavigne, R. J. and D. S. Dennis. 1975. Ethology of 
Efferia frewingi with a description of the eggs 
(Diptera: Asilidae). Annals of the Entomological 
Society of America 68: 992-996. 

McDade, L. A., K. S. Bawa, H. A. Hespenheide and 
G. S. Hartshorn, eds. 1994. La Selva: Ecology and 
Natural History of a Neotropical Rain Forest. The 
University of Chicago Press, Chicago, 486 pp. 

Mensah, R. K. 1996. Evaluation of coloured sticky 
traps for monitoring populations of Austroasca 
vividigrisea (Paoli) (Hemiptera: Cicadellidae) on 
cotton farms. Australian Journal of Entomology 
35(4): 349-353. 

Scarbrough, A. G. 1982. Coexistence in two species 
of Holcocephala (Diptera: Asilidae) in a Mary- 
land habitat: Predatory behavior. Proceedings of 
the Entomological Society of Washington 84(2): 
349-365. 

Shelly, T. E. 1984a. Prey selection by Atractia mar- 
ginata. Proceedings of the Entomological Society 
of Washington 86(1): 120-126. 

. 1984b. Prey selection by the neotropical spi- 

der Micrathena schreibersi with notes on web-site 

tenacity. Proceedings of the Entomological Soci- 

ety of Washington 86(3): 493-502. 

. 1988. Relative abundance of day-flying in- 
sects in treefall gaps vs. shaded understory in a 
neotropical forest. Biotropica 20(2): 114-119. 

Weeks, L. and H. A. Hespenheide. 1985. Predatory 
and mating behavior of Stichopogon (Diptera: As- 
ilidae) in Arizona. Pan-Pacific Entomologist 
61(2): 95-104. 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 652-677 


A REVISION OF THE SHORE-FLY GENUS LEMNAPHILA CRESSON 
(DIPTERA: EPHYDRIDAE) 


WAYNE N. MATHIS AND JAMES E EDMISTON 


(WNM) Department of Entomology, National Museum of Natural History, Smithsonian 
Institution, Washington, DC 20560-0169, U.S.A. (e-mail: mathis.wayne@nmnh.si.edu); 
(JFE) Director, Institute for Franciscan Environmental Studies, Quincy University, Quincy, 
IL 62301, U.S.A. (e-mail: edmisja@quincy.edu) 


Abstract.—Species of the shore-fly genus Lemnaphila Cresson are revised, including 
the description of two new species: L. grossoae (Argentina. Tucuman: San Miguel de 
Tucuman) and L. longicera (Peru. Loreto: Iquitos). Lemnaphila is known thus far only 
from the New World, where there are now five species. All species with known biologies 
are miners in various species of Lemnaceae. Although Lemnaphila is clearly a monophy- 
letic clade based on several synapomorphies, its relationship to other genera of the tribe 
Hydrelliini is unresolved. 


Key Words: Revision, Diptera, Ephydridae, shore flies, Lemnaphila, L. longicera, L. gros- 


soae, New World 


Specimens of Lemnaphila Cresson are 
among the smallest shore flies, with adult 
flies often being less than one mm in length. 
Their small size corresponds well with, and 
is apparently an adaptation to, the mining 
behavior of the larval instars in the tiny 
thalli of the duckweed family Lemnaceae, 
especially the genus Lemna L. Their small 
size evidently has been a deterrent to their 
study, as specimens are seldom collected 
and are poorly represented in collections, 
and very few studies that concern the genus 
have been published. Duckweed, moreover, 
is often an overlooked component of aquat- 
ic environments, and studies pertaining to 
the rearing and identification of phytopha- 
gous insects associated with species of 
Lemnaceae are uncommon (Scotland 1940, 
Buckingham 1989). Our experience in the 
field, however, indicates that specimens of 
Lemnaphila are frequently numerous when 
the duckweed habitat is adequately sam- 


pled. Indeed, recent collecting has revealed 
a fifth species from Peru, a sixth species 
from Argentina, and also major extensions 
in the distributions of the other species. Ad- 
ditional studies on their natural history, in- 
cluding fresh larval and puparial material of 
L. scotlandae Cresson, were conducted in 
Ohio. Describing and documenting these 
discoveries are the primary objectives for 
this revision. Further information on the 
natural history of L. scotlandae will be pub- 
lished separately (Edmiston and Foote, in 
preparation). 

Cresson (1933) first described Lemna- 
phila as a monotypic genus, being known 
only from L. scotlandae Cresson. Miss 
Minnie B. Scotland, after whom this spe- 
cies was named, discovered the rather mi- 
nute adults of L. scotlandae on Lemna that 
was growing in a fish hatchery in Ithaca, 
New York. Miss Scotland conducted studies 
on this species and published three papers 


VOLUME 102, NUMBER 3 


(1934, 1939, 1940) on its natural history. 
The genus remained monotypic until Lizar- 
ralde de Grosso (1977, 1978) described 
three additional species, all based on spec- 
imens collected in the neotropics. In the 
first paper, Lizarralde de Grosso (1977) de- 
scribed two of the new species: L. neotro- 
pica (Argentina (La Plata), Panama, and Ja- 
maica) and L. wirthi (Ecuador). Lizarralde 
de Grosso (1978) described the third spe- 
cies, L. lilloana, from specimens collected 
in Argentina (Tucuman). In the second pa- 
per, she also described the immature stages 
of L. lilloana and L. neotropica and report- 
ed the occurrence of L. wirthi in Argentina 
(Tucuman). Lizarralde de Grosso (1980) 
published a third paper in which the larvae 
of Lemnaphila are distinguished from other 
shore flies. 

Other than catalog entries (Wirth 1965, 
Mathis and Zatwarnicki 1995), new state 
records (Deonier and Regensburg 1978, 
Steinly et al. 1987), and records of parasa- 
toids (Marsh 1979, Muesebeck 1979), few 
published records on Lemnaphila are avail- 
able. Johannsen (1935) included Lemnaphi- 
la in his generic key to the immature stages 
of shore flies, as did Courtney et al. (1996) 
and Merritt et al. (1996). The latter two pa- 
pers also summarized known information 
on the biology of the larvae. Wirth and 
Stone (1956) and Wirth et al. (1987) in- 
cluded Lemnaphila in their keys to the 
shore-fly genera of the Nearctic Region. In 
a synopsis of shore-fly biology, Foote 
(1995) also reviewed the biology of Lem- 
naphila. 

The descriptive terminology, with the ex- 
ceptions noted in Mathis (1986) and Mathis 
and Zatwarnicki (1990a), follows that pub- 
lished in the Manual of Nearctic Diptera 
(McAlpine 1981). Because specimens are 
small, usually less than 3.5 mm in length, 
study and illustration of the male terminalia 
required use of a compound microscope. 
Although we followed the terminology for 
most structures of the male terminalia that 
other workers in Ephydridae have used (see 
references in Mathis 1986; Mathis and Za- 


653 


twarnicki 1990a, 1990b), Zatwarnicki 
(1996) now uses alternative terms (medan- 
drium, transandrium) that are based on the 
‘“‘hinge”’ hypothesis for the origin of the er- 
emoneuran hypopygium. The terminology 
for structures of the male terminalia is pro- 
vided directly on Figs. 4—6 and is not re- 
peated for comparable illustrations of other 
species. 

Dissections of male genitalia were per- 
formed using the method of Clausen and 
Cook (1971) and Grimaldi (1987). External 
morphology and internal genitalic features 
were drawn using a drawing tube on a Wild 
M20EB compound and a MS dissecting mi- 
croscope. The species descriptions are com- 
posite and are not based solely on the ho- 
lotypes. Two head and two venational ratios 
used in the descriptions are defined below 
(all ratios are based on measurements from 
a minimum of five specimens). All mea- 
surements were made from pinned speci- 
mens. 

Eye ratio is the eye width divided by the 
eye height. 

Gena-to-eye ratio is the genal height 
measured at the maximum eye height di- 
vided by the eye height. 

Costal vein ratio is the straight line dis- 
tance between the apices of R,,; and R,,./ 
distance between the apices of R, and R,,;. 

M vein ratio is the straight line distance 
along vein M between crossveins (dm-cu 
and r-m)/distance apicad of dm-cu. 

The phylogenetic analysis was performed 
with the assistance of Hennig86©, a com- 
puterized algorithm that produces clado- 
grams by parsimony. Before analysis, char- 
acter data were arranged in transformation 
series and then polarized, primarily using 
outgroup procedures. 

Rearing.—All field and laboratory work 
was carried out in northeastern Ohio near 
the city of Kent in Portage County. Labo- 
ratory rearings, providing life cycle data, 
were carried out in an environmental cham- 
ber programmed to give a photoperiod of 
16L:8D and a temperature variation be- 
tween 22-—32° C. Adults were collected 


654 


from a marsh near the Kent State University 
stadium and placed in plastic tumblers with 
approximately 100 ml of pond water. Lem- 
na thalli (approximately 25) were placed in 
each tumbler. The tumbler was covered 
with cheese cloth held in place by a rubber 
band. 

Immature descriptions.—Specimens col- 
lected from the field or reared in the labo- 
ratory were preserved in 70% ethanol or 
double mounted and dried. External and in- 
ternal morphologic structures were drawn 
using an ocular grid on a Zeiss compound 
microscope. Continuous characters were 
measured using a calibrated ocular micro- 
meter attached to a Zeiss compound micro- 
scope. Descriptive terminology of immature 
stages follows McAlpine (1981). The ten- 
toropharyngeal sclerite length and width are 
the maximum straight line distances in lat- 
eral view from the anterior tip to the dorsal 
cornu posterior tip (A-PDC), from the an- 
terior tip to the ventral cornu posterior tip 
(A-VDC), and from the dorsal cornu pos- 
terior tip to the ventral cornu posterior tip 
(PDC-VDC). 

Fecundity.—Females collected from the 
field were isolated in individual tumblers. 
The number of eggs laid on Lemna thalli 
were counted on each of four days. Each 
day, the female was removed from the tum- 
bler and placed in a fresh preparation with 
25 new Lemna thalli. 

Life History.—Eggs deposited on the 
Lemna thalli were placed individually in 20 
ml plastic vials. Eggs were observed each 
day for hatching, and individual flies were 
observed each day for molting between in- 
stars, pupariation, and adult emergence. 

Specimens.—Although most specimens 
for this study are in the National Museum 
of Natural History, Smithsonian Institution, 
Washington, D.C. (USNM), we also studied 
numerous specimens that are deposited in 
the American Museum of Natural History 
(AMNH), New York; the Academy of Nat- 
ural Sciences of Philadelphia (ANSP), 
Pennsylvania; The Natural History Muse- 
um (BMNH), London, England; Canadian 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


National Collection (CNC), Ottawa, Ontar- 
io; Cornell University (CU), Ithaca, New 
York; Guelph University (GUE), Guelph, 
Ontario; personal collection of James FE Ed- 
miston (JEPC); Instituto Miguel Lillo 
(IML), Tucuman, Argentina; Ohio State 
University (OHSU), Columbus; Universi- 
dad de La Plata, La Plata, Argentina 
(UNLP) and Washington State University 
(WSU). 


SYSTEMATICS 


We follow Deonier’s (1995) tribal char- 
acterization of Hydrelliini and the generic 
composition of the tribe that Mathis and Za- 
twarnicki (1995) published. Deonier (1995) 
published a key to the three genera, includ- 
ing Cavatorella, which he described in the 
same paper. Deonier’s concept of Lemna- 
phila, as reflected in the key, was based pri- 
marily on L scotlandae, the type species, 
but with the addition of other species, the 
characterization of the genus has been ad- 
justed as indicated in the following revised 
key to genera. 


KkyY TO GENERA OF HYDRELLIINI 
ROBINEAU-DESVOIDY 


(modified from Deonier 1995) 


1. Posterior notopleural seta inserted near dorsal 
angle; prescutellar acrostichal seta lacking; 
ocellar setae lacking; anepisternum bearing a 
single seta along posterior margin; lacking an 
intrapostalar setula ....... Lemnaphila Cresson 

— Posterior notopleural seta inserted at same lev- 

el as anterior seta near ventral margin; pres- 
cutellar acrostichal seta present; ocellar setae 
usually present; anepisternum usually bearing 
1 large seta and 1-2 smaller setae along pos- 
terior margin; usually bearing 1 intrapostalar 
setula 
. Wing apex broadly rounded; vein R,,; long, 
making costal section II much longer than sec- 
tion III; interfacetal setulae conspicuous and 
dense; male lacking ventroclinate processes 
from dorsad of fused surstyli............. 

Hydrellia Robineau-Desvoidy 

— Wing apex narrowly rounded, ellipsoidal; vein 
R,,; short, making costal section II subequal 
to section III; costa bearing several interfrac- 
tural setae; interfacetal setulae inconspicuous, 
sparse; male with ventroclinate styliform pro- 


Nw 


VOLUME 102, NUMBER 3 


cesses projected from dorsad of fused surstyli 
Peay See hs Sh sO ecient Cavatorella Deonier 


Genus Lemnaphila Cresson 


Lemnaphila Cresson, 1933:229. Type spe- 
cies: Lemnaphila scotlandae Cresson, 
1933, monotypy; 1944:175—176 [review, 
tribal relationships].-Scotland, 1934:291 
[biology, larval miner in Lemna]; 1939: 
713-718 [biology, parasatoids]; 1940: 
322-324 [review of biology, figs. of egg, 
puparium, adults].—Johanssen, 1935:48— 
51 [larval key].—Wirth, 1965:745 [Ne- 
arctic catalog].—Wirth and Stone, 1956: 
465, 469 [key to genus, leaf mining hab- 
it]|—Lizarralde de Grosso, 1977:159-— 
164 [review]; 1980:55—60 [larval key]; 
1989:54—55 [review, Argentina fauna]. 
Wirth et al., 1987:1035, 1041 [fig. of 
head, key to genus].—Mathis and Za- 
twarnicki, 1995:96—97 [world catalog].— 
Courtney et al., 1996:508 [larval key to 
genus].—Merritt et al., 1996:545 [habitat, 
habits, trophic relationships]. 


Diagnosis.—Lemnaphila is a genus of 
the tribe Hydrelliini, subfamily Hydrelli- 
inae, that is similar and closely related to 
Hydrellia Robineau-Desvoidy and Cavato- 
rella. Lemnaphila, which are minute to 
small shore flies (length 0.83—1.50 mm), is 
distinguished from these and other shore-fly 
genera by the following combination of 
characters: 

Head: Frons much wider than long, 
sometimes with distinct mesofrons and par- 
afrons; mesofrons broadly reaching anterior 
margin of frons, vestiture variable. Ocellar 


and paravertical setae lacking; pseudopos- | 


tocellar setae well developed, proclinate 
and slightly divergent. Pedicel lacking a 
large, dorsoapical seta; arista with 5—9 long 
rays along dorsum. Face in lateral view ver- 
tically straight or shallowly concave and 
with ventral third slightly angled anteriorly; 
ventral facial margin at most shallowly 
arched; facial seta 1 near ventrolateral mar- 
gin, sometimes with a few setulae, gener- 
ally weakly developed. Eye comparatively 
large, especially in lateral view, bearing nu- 


655 


merous interfacetal setulae. Gena very 
short, gena-to-eye ratio less than 0.15; lack- 
ing a prominent genal seta. Clypeus not 
generally exposed. 

Thorax: Chaetotaxy: dorsocentral setae 2 
(0+2 or with anterior seta on transverse su- 
ture), posterior dorsocentral seta displaced 
laterally (this seta has sometimes been la- 
beled as the intra-alar seta); lacking pres- 
cutellar acrostichal setae, other acrostichal 
setae variable, either lacking or with a few 
setulae in 2 anterior rows; presutural supra- 
alar seta present but often reduced; lacking 
postpronotal, postsutural supra-alar, and in- 
trapostalar setae; 1 postalar seta; 2 lateral 
scutellar setae (lacking a smaller setula be- 
tween 2 larger setae), basal seta shorter than 
apical seta; 2 notopleural setae, anterior seta 
usually shorter, posterior seta inserted near 
dorsal angle of notopleuron; 1 anepisternal 
seta along posterior margin; katepisternal 
seta variable, lacking, reduced, or well de- 
veloped. Wing: generally hyaline or very 
faintly infuscate; costal vein long, extended 
to vein M; vein R,,, short, with costal sec- 
tion II equal to section IJ or much shorter 
than section III; lacking greatly elongate in- 
terfractural setulae other than apical seta. 
Legs lacking dorsoerect setae on dorsal sur- 
face. 

Abdomen: Dorsum comprising tergites 
1-5, with tergites 1-2 somewhat fused; 
sternite 1 greatly reduced or lacking, 2—5 
present; 5th sternite of male comparatively 
well developed and much larger than ster- 
nites 2—4, bearing setulae, shape variable 
with species. Male terminalia: Epandrium 
weakly developed, incomplete dorsally with 
a membranous gap between lateral exten- 
sions, lateral extensions as a fused lateral 
phalange at base of cerci; cerci well devel- 
oped, fused with median margin of epan- 
drium; surstylus greatly reduced as a small, 
usually narrow process from ventral margin 
of fused epandrium/cerci; subepandrial 
plate well developed, simple, arched; ae- 
deagus well developed, sometimes angulate 
in lateral view; aedeagal apodeme narrow 
to very broadly developed, keel variously 


656 


developed, usually enlarged; postgonite 
well developed, usually bearing setulae 
near midlength; pregonite apparently lack- 
ing; hypandrium simple, lateral apices ori- 
ented posteriorly, shallowly arched or nar- 
rowly V-shaped. 

Discussion.—As noted in the generic di- 
agnosis, we place Lemnaphila in the tribe 
Hydrelliini, subfamily Hydrelliinae, where 
it is closely related to Hydrellia and Ca- 
vatorella. Although this tribal assignment 
and the generic associations are now ap- 
parent, these relationships were not always 
recognized and there was some confusion 
that led to a rather circular history. When 
Cresson (1933) described Lemnaphila he 
placed the genus “. . . in the tribe Hydrel- 
liae of the subfamily Notiphilinae, near the 
genus Nostima.’’ Cresson based his generic 
placement on adult characters (setulose 
eyes, antennal shape, and the arrangement 
and number of dorsocentral setae). Cresson 
(1944), however, then removed WNostima 
Coquillett, Philygria Stenhammar, and 
Lemnaphila from the tribe Hydrelliini and 
created a new tribe, Hydrinini (unavailable, 
based on a junior homonym, = Philygriini 
Lizarralde de Grosso), for these three gen- 
era. Despite removing Lemnaphila from a 
close association with Hydrellia, Cresson 
(1944:176) continued to recognize that 
Lemnaphila has ‘‘. . . the appearance of a 
small Hydrellia.”’ The tribe Philygriini was 
then characterized primarily by the elevated 
insertion of the posterior notopleural seta 
and the reduced setation. Wirth (1965) fol- 
lowed Cresson’s suprageneric arrangement 
for the Tribe Philygriini in his catalog of 
North American Ephydridae. Dahl (1959), 
in his review of Scandinavian Ephydridae, 
removed Nostima and Philygria from the 
subfamily Notiphilinae (= Hydrelliinae) 
and placed these genera along with Hyadi- 
na Haliday in the subfamily ‘“‘Hydrininae”’ 
(unavailable, based on a junior homonym, 
= Ilytheinae Cresson), which Dahl de- 
scribed in the same paper. Mathis and Za- 
twarnicki (1990b) also recognized Philygri- 
ini but with only two genera: Philygria and 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Nostima. They suggested that Lemnaphila 
was better placed in the tribe Hydrelliini 
(Subfamily Hydrelliinae) near the genus 
Hydrellia based especially on characters of 
the male terminalia and larvae, and in sub- 
sequent publications (Zatwarnicki 1992, 
Deonier 1995, and Mathis and Zatwarnicki 
1995), Lemnaphila was placed in the Hy- 
drelliini, along with Hydrellia and Cavato- 
rella. 

Cresson (1933, 1944) first noted that 
adults of Lemnaphila are similar to Ay- 
drellia, and a close relationship between 
these two genera is strongly supported by 
larval characters as well. Johannsen (1935) 
was able to distinguish between larvae of 
Lemnaphila and Hydrellia only on the basis 
of larval size and the preference Lemna- 
phila has for feeding on Lemna. Hydrellia 
is generally characterized by its stem- and 
leaf-mining habits, and Johannsen observed 
that the mouthparts of Lemnaphila and Hy- 
drellia were identical except for size. Hen- 
nig (1943) also proposed a close relation- 
ship between larvae of Lemnaphila and Hy- 
drellia. Lizarralde de Grosso (1977, 1978) 
described additional Neotropical species of 
Lemnaphila, including larval descriptions, 
and indicated that the Neotropical larvae of 
Hydrellia and Lemnaphila are very similar 
and closely related. 

Specimens of Lemnaphila are not only 
tiny but they are highly derived, in part and 
presumably as an adaptation to the mining 
habit in Lemnaceae. The structures of the 
male terminalia also reflect their derived 
condition. The surstylus, for example, is 
highly modified as a much reduced struc- 
ture at the ventral margin of the fused epan- 
drium/cercus. The reduced epandrium that 
is fused to the cercus and the enlarged cerci 
are also evidence of the derived status. 


Key TO SPECIES OF LEMNAPHILA CRESSON 


1. Scape, pedicel, and basal half of 1st flagellom- 
ere yellow; length of Ist flagellomere twice its 
height; face in profile straight; length of outer 
vertical seta less than half length of inner ver- 
tical seta. Katepisternum whitish yellow, con- 
colorous with forecoxa 


VOLUME 102, NUMBER 3 


L. longicera, new species 
— Antenna entirely dark brown; length of Ist fla- 
gellomere only slightly longer than its height; 
face shallowly concave in lateral view with 
ventral third slightly angled forward; length of 
outer vertical seta about 3/4 length of inner 
vertical seta. Katepisternum brown .,...... 2 
. Fronto-orbital setulae greatly reduced, at most 
hairlike, none well developed; mesofrons sub- 
shiny, distinct from densely microtomentose 
parafrons. Scutellum lacking microtomentum 
or sparsely microtomentose, not appearing vel- 
vety; katepisternal seta reduced, hairlike ..... 
L. scotlandae Cresson 
— At least | fronto-orbital seta well developed, 
anterior setula proclinate, posterior seta much 
longer, length subequal to outer vertical seta, 
lateroclinate, parallel to outer vertical seta; me- 
sofrons and parafrons densely microtomentose, 
appearing velvety, or with small area immedi- 
ately anterior of ocellar triangle bare (L. /il- 
loana), shiny. Scutellum moderately to densely 
microtomentose, brownish black to black, 
sometimes appearing velvety; katepisternal 
seta well developed, length subequal to poste- 
MOmAnepIStCHnaleScla weer een eka eMenen ens one 3 
3. Ventral third of face densely microtomentose, 
blackish brown, concolorous with dorsal por- 
tion of face; coxae, femora, and tibiae yellow 
PCR e ee, tier L. wirthi Lizarralde de Grosso 
— Ventral third of face densely microtomentose, 
silvery white, distinctly contrasted with black- 
ish brown dorsal portion; femora brown, oth- 
erwise legs yellow 
4. Area immediately laterad and anteriad of ocel- 
lar triangle silvery white, concolorous with 
fronto-orbit immediately anterior and posterior 
of lateroclinate fronto-orbital seta .......... 
L. grossoae, new species 
— Area immediately laterad and anteriad of ocel- 
lar triangle dark brown, similar to coloration of 
OcCellaratian leew ccs. HHS ee eects & ele 5 
5. Mesofrons and parafrons densely microtomen- 
tose, appearing velvety blackish brown; pos- 
terior fronto-orbit extensively white to silvery 
Witte 5 6 G6 bE L. neotropica Lizarralde de Grosso 
— Mesofrons just anterior of ocellar triangle bare, 
shiny, blackish brown, contrasted with densely 
microtomentose parafrons; posterior fronto-or- 
bit with very small area white ............ 
L. lilloana Lizarralde de Grosso 


Nw 


Lemnaphila longicera Mathis and 
Edmiston, new species 
(Figs. 1-3) 


Diagnosis.—This species is distinguished 
from congeners by the following combina- 


657 


tion of characters: Scape, pedicel, and basal 
half of Ist flagellomere yellow; length of 
Ist flagellomere twice height; face in profile 
straight; face mostly uniformly microto- 
mentose and colored, coloration varies de- 
pending on angle of view, except for nar- 
row, vertical stripe; length of outer vertical 
seta less than half inner vertical seta; scu- 
tellum densely microtomentose, appearing 
velvety, dark brown; katepisternum whitish 
yellow, concolorous with forecoxa; legs en- 
tirely yellow, with coxae whitish yellow 
and apical half of tibiae slightly brownish 
yellow. 

Description.—Minute to small shore 
flies, length 0.90—1.35 mm. 

Head: Frons with distinct, subshiny me- 
sofrons; parafrons densely microtomentose, 
dark brown, appearing velvety. Fronto-or- 
bital setae very weakly developed and/or 
lacking, at most with 1-2 hairlike setulae; 
length of outer vertical seta less than half 
inner vertical seta, outer seta curved later- 
ally, inner seta very shallowly curved, near- 
ly vertical; ocelli arranged in equilateral tri- 
angle, or nearly so. Scape, pedicel, and bas- 
al half of Ist flagellomere yellow, apical 
half of Ist flagellomere brown; length of Ist 
flagellomere twice height, apex bluntly 
rounded; arista with 8—9 long, dorsal rays. 
Face in profile straight at anterior margin; 
varying in color from different angles, from 
lateral view mostly yellowish with brown- 
ish yellow area at midheight, from anterior 
view whitish gray, sometimes slightly dark- 
er medially at midheight, and with keel-like 
brown vertical stripe. Eye ratio 0.68—0.71. 
Gena-to-eye ratio 0.10—0.13. Maxillary pal- 
pus yellow; mouthparts mostly yellow to 
grayish yellow. 

Thorax: Mostly dark brown; scutum dark 
brown with moderately dense whitish gray 
microtomentum; scutellum with distal % 
densely microtomentose, dark brown, ap- 
pearing velvety, basal % whitish gray; no- 
topleuron, area of scutum immediately pos- 
teriad of notopleuron densely microtomen- 
tose, dark brown, appearing velvety; anepis- 
ternum and anepimeron concolorous, dark 


658 


Figs. 1-3. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Lemnaphila longicera. 1, Male terminalia (epandrium, cercus, surstylus), lateral view. 2, Internal 


male terminalia (aedeagus, aedeagal apodeme, postgonite, hypandrium, and subepandrial plate), ventral view. 3, 


Fifth sternite, ventral view. 


brown but lighter than notopleuron; katepis- 
ternum, meron, and coxae concolorous, 
whitish yellow. Chaetotaxy: anterior dorso- 
central seta inserted near level of transverse 
suture; acrostichal setae lacking; basal scu- 
tellar seta about % length of apical seta; ka- 
tepisternal seta lacking. Wing: length 1.08— 
1.20 mm; width 0.45—0.49 mm; vein R,,; 
shallowly sinuous, shallowly curved toward 
posterior margin, than shallowly recurved, 


bowed anteriorly; costal vein ratio 1.0—1.07, 
length of costal section II about equal to sec- 
tion III; M vein ratio 0.42—0.47. Halter knob 
bright yellow; stem brownish yellow. Legs: 
femora whitish yellow to yellow basally, be- 
coming more yellow apically; tibiae yellow, 
becoming slightly darker apically; foretarsi 
brownish yellow to brown, mid- and hind- 
tarsus slightly lighter than foretarsus, more 
yellowish. 


VOLUME 102, NUMBER 3 


Abdomen: Dark brown, male Sth tergite 
subshiny; 5th sternite of male (Fig. 3) well 
developed, narrowly heart shaped, longer 
than wide, tapered gradually to narrowly 
rounded anterior margin, posterior margin 
deeply emarginate. Male terminalia (Figs. 
1—2): Epandrium (Fig. 1) reduced to nar- 
row, fused, lateral phalange at basolateral 
margin of cercus; cercus (Fig. 1) greatly en- 
larged, projected posteriorly, bearing 4—5 
large setulae at anteroventral margin; su- 
bepandrial plate simple, arched, connected 
ventrally with postgonite; aedeagus in lat- 
eral view (Fig. 1) with moderately broad 
base tapered irregularly thereafter to narrow 
process, apex narrowly rounded; aedeagal 
apodeme with ventral keel comparatively 
long and narrow in lateral view (Fig. 1), 
slightly asymmetrical; postgonite prominent 
with rounded apex, bearing setulae about %4 
distance from connection with subepandrial 
plate; hypandrium V-shaped in ventral view 
(Fig. 2): 

Type Material—The holotype male is la- 
beled ““PERU. Loreto: Iquitos [,] 16Feb1984 
[,] Wayne N. Mathis/sHOLOTYPE Lemna- 
phila longicera 6 W.N. Mathis & Edmiston 
USNM [red; species name, gender, and au- 
thors handwritten].’” The holotype is double 
mounted (minuten in a block of plastic), is 
in excellent condition, and is deposited in 
the USNM. Twenty-two paratypes (8 d, 14 
2; USNM) bear the same label data as the 
holotype. 

Distribution.—WNeotropical: Peru (Lore- 
to). 

Etymology.—The species epithet, longi- 
cera, alludes to the elongate antenna that, 
among other characters, distinguishes this 
species. 

Remarks.—Although this species clear- 
ly belongs in Lemnaphila, it is very dis- 
tinct structurally (see diagnosis and key) 
and apparently represents a lineage that 
has undergone considerable anagenetic 
evolution. We know nothing about the nat- 
ural history of the immature stages but 
suggest that they will be found to be min- 
ers in Lemnaceae. 


659 


Lemnaphila grossoae Mathis and 
Edmiston, new species 


Diagnosis.—This species is distinguished 
from congeners by the following combina- 
tion of characters: Antenna entirely dark 
brown; length of Ist flagellomere only 
slightly longer than height; face in lateral 
view shallowly concave; ventral third of 
face densely microtomentose, silvery white, 
distinctly contrasted with blackish brown 
dorsal portion; mesofrons densely microto- 
mentose, area immediately laterad and an- 
teriad of coellar triangle silvery white, oth- 
erwise brown to blackish brown; fronto-or- 
bit anteriad and posteriad of long laterocli- 
nate fronto-orbital seta silvery white, 
otherwise velvety blackish brown; scutel- 
lum sparsely microtomentose, appearing 
subshiny to faintly dull, brown; katepister- 
num brown; femora brown; tibiae and most 
tarsomeres yellow. 

Description.—Small shore flies, length 
1.30 mm. 

Head: Frons entirely microtomentose, bi- 
colored, brown and silvery white; parafrons 
and anterior half of fronto-orbit concolo- 
rous, densely microtomentose, brownish 
black, appearing velvety, area immediately 
laterad and anteriad of ocellar triangle sil- 
very white; posterior fronto-orbit (from 
base of outer vertical seta to base of later- 
oclinate fronto-orbital seta) with silvery 
white microtomentum, concolorous with 
area laterad and anteriad of ocellar triangle. 
One fronto-orbital seta well developed, 
length subequal to outer vertical seta, ori- 
entation postero-obliquely lateroclinate to 
lateroclinate; other fronto-orbital setae an- 
terior of large seta, at best weakly devel- 
oped, hairlike; length of outer vertical seta 
about *% that of inner vertical seta, outer 
vertical seta distinctly curved laterally, in- 
ner vertical seta very shallowly curved, 
nearly vertical; ocelli arranged in isosceles 
triangle, with distance between posterior 
ocelli greater than between either posterior 
ocellus and anterior ocellus. Antenna en- 
tirely dark brown; length of Ist flagellom- 


660 


ere only slightly longer than height; arista 
with 6—7 long, dorsal rays. Face shallowly 
concave in lateral view; ventral third of face 
densely microtomentose, silvery white, dis- 
tinctly contrasted with blackish brown dor- 
sal portion; dorsal portion of face with very 
shallow carina, mostly faintly grayish black 
except for black area on either side of mid- 
line just dorsad of silvery white microto- 
mentum. Eye ratio 0.70. Gena-to-eye ratio 
0.19. Maxillary palpus blackish brown; 
mouthparts generally brownish black. 

Thorax: Mostly blackish brown; scutum 
subshiny, sparsely microtomentose; scutel- 
lum with moderately dense microtomen- 
tum, brownish black, somewhat dull, not 
velvety appearing like frons except partially 
along sides; pleuron mostly blackish brown 
to black except most of anepisternum ap- 
pearing silvery gray to silvery white. Chae- 
totaxy: anterior dorsocentral seta inserted 
behind level of transverse suture; acrosti- 
chal setulae sparse anteriorly, in 2 rows, 
lacking posteriorly; basal scutellar seta 
about % length of apical seta; katepisternal 
seta well developed, length subequal to pos- 
terior anepisternal seta. Wing: length 1.30 
mm; width 0.50 mm; vein R,,; shallowly 
bowed anteriorly; costal vein ratio 1.25; 
length of costal section II about half of sec- 
tion III; M vein ratio 0.55. Halter knob 
bright yellow; stem blackish brown. Legs: 
Femora mostly brown, apices yellowish; 
tibiae and most tarsomeres yellow, 5th tar- 
somere brown. 

Abdomen: Tergites very sparsely micro- 
tomentose to bare, blackish brown. 

Type Material.—The holotype female is 
labeled ““ARGENTINA San Miguel de 
TUCUMAN 2-XII-1976 Col. L. de Grosso 
[handwritten]/Lemnaphila lillolana det. M. 
Lizarralde de Grosso/PARATIPO [yellow]/ 
COLECCION INST.-FUND M. LILLO 
(4000)—S.M. TUCUMAN TUCUMAN— 
ARGENTINA [yellow]/HOLOTYPE Lem- 
naphila grossoae 2 W.N. Mathis & Edmis- 
ton USNM [red; species name, gender, and 
authors handwritten].’’ The holotype is 
double mounted (glued to a large triangle), 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


is in good condition (glue partially obscur- 
ing tarsi), and is deposited in the IML. 

Distribution.—WNeotropical: Argentina 
(Tucuman). 

Etymology.—The species epithet, gros- 
soae, is a patronym to honor and recognize 
the numerous contributions of Dr. Mercedes 
Lizarralde de Grosso to the study of shore 
flies, Lemnaphila in particular. 

Remarks.—This species was misidenti- 
fied as L. lillolana but the pattern of micro- 
tomentum and coloration of the head, es- 
pecially the frons, differs from that species 
as noted in the diagnosis. In addition to the 
holotype female, we dissected the male al- 
lotype/paratype of L. lillolana and discov- 
ered that it is likewise not conspecific with 
L. lillolana. Our basis for this conclusion is 
comparison of the structures of the male 
terminalia with those that Lizarralde de 
Grosso (1978:15, figs. 6-13) published. 
Thus, this male, which lacks its head, may 
be conspecific with this species. 


Lemnaphila lillolana Lizarralde de Grosso 


Lemnaphila lillolana Lizarralde de Grosso, 
1978:14-17 [immature stages; host 
plants: Spirodela biperforata Kock, Wol- 
fiella lingulata (Hegelm.), Salvinia mini- 
ma Baker (Lemnaceae)]; 1989:55  [re- 
view, Argentina fauna].-Mathis and Za- 
twarnicki, 1995:96 [world catalog]. 


Diagnosis.—This species is distinguished 
from congeners by the following combina- 
tion of characters: Antenna entirely dark 
brown; length of Ist flagellomere only 
slightly longer than height; face in lateral 
view shallowly concave; ventral third of 
face densely microtomentose, silvery white, 
distinctly contrasted with blackish brown 
dorsal portion; mesofrons just anterior of 
ocellar triangle bare, shiny, blackish brown, 
contrasted distinctly with densely microto- 
mentose parafrons; posterior fronto-orbit 
with very small area white; scutellum 
densely microtomentose, appearing velvety, 
dark brown; katepisternum brown; femora 
brown; tibiae and most tarsomeres yellow. 


VOLUME 102, NUMBER 3 


Description.—Small shore flies, length 
1.40—1.50 mm. 

Head: Frons with mesofrons except for 
ocellar triangle bare of microtomentum, 
shiny, blackish brown; parafrons, and an- 
terior half of fronto-orbit concolorous, 
densely microtomentose, brownish black, 
appearing velvety; posterior fronto-orbit 
(from base of outer vertical seta to base of 
lateroclinate fronto-orbital seta) with whit- 
ish gray microtomentum. One fronto-orbital 
seta well developed, length subequal to out- 
er vertical seta, orientation postero-oblique- 
ly lateroclinate to lateroclinate; other fron- 
to-orbital setae anterior of large seta, at best 
weakly developed, hairlike; length of outer 
vertical seta about % that of inner vertical 
seta, outer vertical seta distinctly curved 
laterally, inner vertical seta very shallowly 
curved, nearly vertical; ocelli arranged in 
isosceles triangle, with distance between 
posterior ocelli greater than between either 
posterior ocellus and anterior ocellus. An- 
tenna entirely dark brown; length of 1st fla- 
gellomere only slightly longer than height; 
arista with 7—8 long, dorsal rays. Face shal- 
lowly concave in lateral view; ventral third 
of face densely microtomentose, silvery 
white, distinctly contrasted with blackish 
brown dorsal portion; dorsal portion of face 
with very shallow carina, mostly faintly 
grayish black except for black area on ei- 
ther side of midline just dorsad of silvery 
white microtomentum. Eye ratio 0.75—0.77. 
Gena-to-eye ratio 0.1 1—0.12. Maxillary pal- 
pus blackish brown; mouthparts generally 
brownish black. 

Thorax: Mostly blackish brown; scutum 
subshiny, sparsely microtomentose; scutel- 
lum with moderately dense microtomen- 
tum, brownish black, dull, not velvety ap- 
pearing like frons; pleuron mostly blackish 
brown to black except dorsal half of ane- 
pisternum from some angles appearing sil- 
very gray. Chaetotaxy: anterior dorsocen- 
tral seta inserted behind level of transverse 
suture; acrostichal setulae sparse anteriorly, 
in 2 rows, lacking posteriorly; basal scutel- 
lar seta about % length of apical seta; ka- 


661 


tepisternal seta well developed, length sub- 
equal to posterior anepisternal seta. Wing: 
length 1.45-1.60 mm; width 0.60—0.74 
mm; vein R,,; shallowly bowed anteriorly; 
costal vein ratio 1.30—1.33; length of costal 
section II about half of section III; M vein 
ratio 0.49—0.50. Halter knob bright yellow; 
stem blackish brown. Legs: Femora mostly 
brown, apices yellowish; tibiae and most 
tarsomeres yellow, 5th tarsomere brown. 

Abdomen: Dark brown. 

Type Material.—The holotype female 
(not a male as indicated in the original de- 
scription) is labeled “‘ARGENTINA[.] San 
Miguel de TUCUMAN[,] 2.XII.1976 [2 
Dec 1976] Col. L. de Grosso [handwritten, 
black margin]/Lemnaphila Lilloana Lizar- 
ralde de Grosso det. Lizarralde de Grosso 
[handwritten, black marginJ/HOLOTIPO 
[red]J/COLECCION INST. - FUND. M. 
LILLO (4000) - S. M. TUCUMAN TUC- 
UMAN - ARGENTINA [yellow].”’ The ho- 
lotype is double mounted (glued to mod- 
erately large paper triangle), is in fair con- 
dition (specimen slightly teneral, eyes and 
face partially collapsed, right wing with 
posterior margin folded back on itself), and 
is deposited in the IML. 

Other Specimens Examined.—ARGEN- 
TINA. Tucumdn: San Miguel de Tucuman, 
2 Dec 1976, M. Lizarralde de Grosso (6 9; 
paratypes; IML). 

Distribution.—Neotropical: Argentina 
(Tucuman). 

Remarks.—This species is structurally 
very similar to L. neotropica and is obvi- 
ously closely related. The only difference 
that we have discovered in external struc- 
tures is the shiny mesofrons with the ex- 
ception of the ocellar triangle (L. neotro- 
pica has a completely microtomentose, vel- 
vety, blackish brown mesofrons). The other 
differences that Lizarralde de Grosso noted 
(1978:17) are variable and are within the 
ranges of either species. No males of this 
species were available for description and 
illustration. 


662 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 4-7. 


Lemnaphila neotropica. 4, Male terminalia (epandrium, cercus, surstylus, subepandrial plate, 


postgonite), posterior view. 5, Same, lateral view. 6, Internal male terminalia, lateral view. 7, Fifth sternite and 


hypandrium, ventral view. 


Lemnaphila neotropica Lizarralde de 
Grosso 
(Figs. 4—7) 


Lemnaphila neotropica Lizarralde de Gros- 
so, 1977:160; 1978:17-18 [biology, im- 
mature stages, host plants: Lemna minima 
Philippi, Hydromystia stolonifera,; para- 
site: Hexacola sp. (Cynipidae)]; 1989:55 
[review, Argentina fauna].—Mathis and 
Zatwarnicki, 1995:96—97 [world catalog]. 


Diagnosis.—This species is distinguished 
from congeners by the following combina- 
tion of characters: Antenna entirely dark 
brown; length of Ist flagellomere only 
slightly longer than height; face in lateral 
view shallowly concave; ventral third of 
face densely microtomentose, silvery white, 


distinctly contrasted with blackish brown 
dorsal portion; mesofrons and parafrons en- 
tirely densely microtomentose, appearing 
velvety blackish brown; scutellum densely 
microtomentose, appearing velvety, dark 
brown; katepisternum brown; femora 
brown; tibiae and most tarsomeres yellow. 

Description.—Minute to small shore 
flies, length 0.90—1.30 mm. 

Head: Frons with mesofrons, parafrons, 
and anterior half of fronto-orbit concolo- 
rous, densely microtomentose, brownish 
black, appearing velvety; posterior fronto- 
orbit (from base of outer vertical seta to 
base of lateroclinate fronto-orbital seta) 
with whitish gray microtomentum. One 
fronto-orbital seta well developed, length 
subequal to outer vertical seta, orientation 


VOLUME 102, NUMBER 3 


postero-obliquely lateroclinate to laterocli- 
nate; other fronto-orbital setae anterior of 
large seta, at best weakly developed, hair- 
like; length of outer vertical seta about % 
that of inner vertical seta, outer vertical seta 
distinctly curved laterally, inner vertical 
seta very shallowly curved, nearly vertical; 
ocelli arranged in isosceles triangle, with 
distance between posterior ocelli greater 
than between either posterior ocellus and 
anterior ocellus. Antenna entirely dark 
brown; length of Ist flagellomere only 
slightly longer than height; arista with 6—7 
long, dorsal rays. Face shallowly concave 
in lateral view; ventral third of face densely 
microtomentose, silvery white, distinctly 
contrasted with blackish brown dorsal por- 
tion; dorsal portion of face with very shal- 
low carina, mostly faintly grayish black ex- 
cept for black area on either side of midline 
just dorsad of silvery white microtomen- 
tum. Eye ratio 0.75—0.77. Gena-to-eye ratio 
0.11—0.12. Maxillary palpus blackish 
brown; mouthparts generally brownish 
black. 

Thorax: Mostly brownish black to black; 
scutum subshiny, sparsely microtomentose; 
scutellum with moderately dense microto- 
mentum, brownish black, dull, not velvety 
appearing like frons; pleuron mostly 
brownish black to black except dorsal half 
of anepisternum from some angles appear- 
ing silvery gray. Chaetotaxy: anterior dor- 
socentral seta inserted behind level of trans- 
verse suture; acrostichal setulae sparse an- 
teriorly, in 2 rows, lacking posteriorly; bas- 
al scutellar seta about % length of apical 
seta; katepisternal seta well developed, 
length subequal to posterior anepisternal 
seta. Wing: length 0.80—1.30 mm; width 
0.45—0.65 mm; vein R,,; shallowly bowed 
anteriorly; costal vein ratio 1.60—1.69; 
length of costal section II slightly less than 
half of section III; M vein ratio 0.46—0.55. 
Halter knob bright yellow; stem blackish 
brown. Legs: Femora mostly brown, apices 
yellowish; tibiae and most tarsomeres yel- 
low, 5th tarsomere brown. 

Abdomen: Dark brown, male 5th tergite 


663 


subshiny; 5th sternite of male in ventral 
view (Fig. 7) much wider than long, broad- 
ly and shallowly U-shaped, bearing numer- 
ous setulae along posterior margin, these 
also in membrane between sternite and hy- 
pandrium. Male terminalia (Figs. 4—5): 
Epandrium comparatively well developed 
as a wide phalange at base of cerci; cerci 
(Figs. 4—5) well developed, irregularly 
hemispherical around posteromedial mar- 
gin, narrowed at dorsal and ventral angles, 
lateral margin nearly straight in lateral 
view, shallowly concave in posterior view; 
surstylus a short process, only slightly lon- 
ger than wide, bearing numerous setulae, 
these longer apically; subepandrial plate 
bandlike in lateral view (Fig. 6), distinctly 
arched in posterior view (Fig. 4); aedeagus 
very well developed, angulate in lateral 
view, robust, especially basiphallus; aedea- 
gal apodeme greatly enlarged, especially 
expanded keel, keel slightly asymmetrical 
in lateral view, fanlike; postgonite promi- 
nent, heavily sclerotized, base in ventral 
view rectangular with a narrowed, apical, 
arched process extended from ventromedial 
portion of base, apical process parallel sid- 
ed, apex narrowly rounded; pregonite ap- 
parently lacking; hypandrium (Fig. 7) in 
ventral view a broad, arched band, posterior 
Margin concave, anterior margin convex, 
with a short, tonguelike, medial process. 

Type Material.—The holotype male is la- 
beled “‘[Argentina. Buenas Aires:] LA 
PLATA 18/10/71 [18 Oct 1971] COL. 
[Loidcono-] DIAZ [handwritten; black sub- 
margin|/HOLOTIPO [red]/$/MUSEO DE 
LA PLATA LEMNAPHILA NEOTROPI- 
CA L. DE GROSSO Det LIZARRALDE 
DE GROSSO [species name and author 
handwritten; black submargin].”’> The ho- 
lotype is double mounted (glued to a large 
paper triangle), is in fair condition (abdo- 
men missing, some setae misoriented), and 
is deposited in the UNLP. 

Other Specimens Examined.—ARGEN- 
TINA. Buenos Aires: La Plata, 18 Oct 1971, 
Loidcono-Diaz (1 6, 1 2; paratypes; 
UNLP). 


664 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 8-10. Lemnaphila scotlandae. 8, Male terminalia (epandrium, cercus, surstylus, aedeagus, aedeagal 
apodeme, subepandrial plate, postgonite), lateral view. 9, Internal male terminalia (surstylus shown on right side 
only, postgonite shown on left side only), dorsal view. 10, Fifth sternite, ventral view. 


DOMINICAN REPUBLIC. La Vega: Jar- 
abacoa (1-2 km S; 19°06.9'N, 70°37'W; 
520 m), 14-17 May 1998, D. and W. N. 
Mathis (11 6, 4 2; USNM). 

GRENADA. St. George: Point Salines 
Airport (W end; 12°00.3'N, 61°47.7'W), 12 
Sep 719979 WiN- Mathis: (7.3659 bh @s 
USNM). 

JAMAICA. St. Ann: Runaway Bay 
(stream bed), Feb 1969, W. W. Wirth (1 od; 
paratype; USNM). 

PANAMA. Aguadulce Cocle, 25 Sep 
1951, ES. Blanton (1 6; paratype, USNM). 

Distribution.—Neotropical: Argentina 
(Buenos Aires, Chaco, Tucuman), Panama, 
West Indies (Dominican Republic, Grenada, 
Jamaica). 

Remarks.—This species and L. wirthi are 
very similar and obviously closely related. 
This relationship is established by the fol- 
lowing synapomorphies: 1. Aedeagus large, 
prominent, angulate; 2. Sth sternite of male 
much wider than long and bearing numer- 
ous setulae along posterior, concave mar- 
gin. 

Lemnaphila scotlandae Cresson 
(Figs. 8-21) 


Lemnaphila scotlandae Cresson, 1933:229 
[USA. New York: Tompkins County, Ith- 


aca, Fall Creek; HT 3, ANSP (6501)]; 
1944:176 [tribal assignment].—Johan- 
nsen, 1935:50—51 [immature stages].— 
Scotland, 1934:291 [biology, larval min- 
er in Lemna]; 1939:713-—717 [host plant: 
Lemna minor L. (Lemnaceae); parasit- 
oids: Opius lemnaphilae Muesebeck 
(Braconidae), Trichopria angustipennis 
Muesebeck, 7. paludis Muesebeck (Dia- 
priidae)]; 1940:322—324 [immature stag- 
es, host plants and parasitoids].—Mue- 
sebeck, 1939:58—62 [descriptions of par- 
asitoids]; 1979:1146—-1147 [catalog of 
parasitoids, Trichopria angustipennis and 
T. paludis|.—Wirth, 1965:745 [Nearctic 
catalog].—Deonier and Regensburg, 
1978:155 [records for Ohio].—Marsh, 
1979:209 [catalog of parasitoid, Opius 
lemnaphilae|.—Ferrar, 1987:169 [sum- 
mary of biology].—Steinly et al., 1987: 
166, 168 [records for Illinois, discus- 
sion].—Buckingham, 1989:219-221 
[parasitoids: Opius lemnaphilae Muese- 
beck (Braconidae), Trichopria angusti- 
pennis Muesebeck, T. paludis Muesebeck 
(Diapriidae); records for Florida; review 
of biology].—Mansor and Buckingham, 
1989:115-118 [host range].—Foote, 
1995:430 [review of biology].—Mathis 


VOLUME 102, NUMBER 3 


and Zatwarnicki, 1995:97 [world cata- 
log]. 


Diagnosis.—This species is distinguished 
from congeners by the following combina- 
tion of characters: Antenna entirely dark 
brown; length of Ist flagellomere only 
slightly longer than height; mesofrons sub- 
shiny, distinct from densely microtomento- 
se parafrons; face in lateral view shallowly 
concave; ventral third of face densely mi- 
crotomentose, appearing velvety, dark 
brown to golden yellow, depending on an- 
gle; scutellum lacking dense microtomen- 
tum, gray, not appearing velvety; katepis- 
ternum brown; legs brown. 

Description.—Minute to very small 
shore flies, length 0.83—1.2 mm; yellowish 
brown, dark brown, to black with silvery 
gray and yellowish silver microtomentum. 

Head: Frons generally moderately dense- 
ly invested with silvery whitish gray micro- 
tomentum; mesofrons from ocellar triangle 
anteriad moderately densely silvery whitish 
gray microtomentose, subshiny; parafrons 
brown; fronto-orbit generally concolorous 
with mesofrons or slightly more brownish. 
Occiput brown with yellowish silver micro- 
tomentum; outer vertical seta lateroclinate; 
length of outer vertical seta about %4 inner 
vertical seta; inner vertical seta parallel to 
slightly convergent; fronto-orbital setae mi- 
nute, hairlike; pseudopostocellar setae 
prominent, proclinate and somewhat diver- 
gent; paravertical setae absent; ocellar setae 
absent; postocullar setae absent; ocelli ar- 
ranged in isosceles triangle, with distance 
between posterior ocelli much greater than 
between either posterior ocellus and ante- 
rior ocellus. Antenna entirely dark brown; 
length of Ist flagellomere only slightly lon- 
ger than its height; arista bearing 5—7 rays 
along dorsum. Facial background colora- 
tion shiny dark brown with sparse silvery 
gray microtomentum. Oral margin slightly 
protruded dorsally. Face with transverse 
crease at about ventral third, shallowly con- 
cave in lateral view, ventral third slightly 
angled forward, densely microtomentose, 


665 


dark brown. Parafacial band shiny dark 
brown. Eye ratio 0.78—0.83. Gena-to-eye 
ratio 0.14—0.17. Gena and postgena brown 
with sparse silvery yellow microtomentum; 
parafacial setae minute; gena bearing small 
black setulae. Maxillary palpus dark brown 
with prominent setae; prementum shiny 
dark brown, bulbous. 

Thorax: Mesonotum brown with silvery 
yellow microtomentum; scutellum lacking 
microtomentum or microtomentum sparse, 
not appearing velvety; scutum length 0.29— 
0.43 mm; scutellar length 0.10—0.17 mm; 
subscutellum brown with yellowish silver 
microtomentum; anepisternum with dorsal 
half brown with dense silvery gray micro- 
tomentum, ventral half with sparse silvery 
and golden microtomentum; katepisternum 
brown with sparse silvery golden microto- 
mentum; anatergite brown with yellowish 
silver microtomentum. Chaetotaxy: anterior 
dorsocentral seta inserted near level of 
transverse suture; acrostichal setulae lack- 
ing; scutellar setae with anterolateral setae 
3%44 length of posteromedial setae and me- 
diolateral setulae; katepisternal seta weakly 
developed, hairlike, much shorter than pos- 
terior anepisternal seta. Wing: length 1.02— 
1.44 mm; width 0.42—0.61 mm; vein R,,; 
shallowly bowed anteriorly; costal vein ra- 
tio 1.76—1.85, with length of costal section 
II about half costal section III; M vein ratio 
0.46—0.55. Halter knob bright yellow; stem 
blackish brown. Legs: yellowish brown to 
brown; tarsomeres 1—4 yellowish brown, 
darker distally; tarsomere 5 brown. 

Abdomen: Tergite 3 length, males 0.13-— 
0.17 mm, females 0.14—0.20 mm; tergite 4 
length, males 0.18—0.20 mm, females 0.18— 
0.20 mm. Background coloration brown to 
dark brown. Tergites 1—5 covered with 
sparse yellow microtomentum; 5th sternite 
slightly longer than wide, tapered slightly 
anteriorly, bearing numerous setulae. Male 
Terminalia (Figs. 8—9): Epandrium much 
reduced, as a small, lateral phalange at base 
of fused cercus; cercus comparatively great- 
ly enlarged, height nearly equal to width of 
5th sternite of male; subepandrial plate an 


666 


Figs. 11-16. 
ventral view. 13, Micropylar end of egg, ventral view. 14, Third-instar larva, cephalic end, lateroblique view. 
15, Same, antennae. 16, Same, posterior end, ventral view. 


arched, narrow, bandlike plate; surstylus 
(Fig. 8) a long, narrow, parallel-sided pro- 
cess bearing 2 apical setulae; aedeagus 
cornucopialike in lateral view (Fig. 8), with 
broad base, tapered to narrow apex that is 
hooked; aedeagal apodeme with keel com- 
paratively long and narrow in lateral view, 
essentially symmetrical; subepandrial plate 
forming rounded arch, connected ventrally 
with postgonite; postgonite (Fig. 8) promi- 
nent with a posterior and anterior process; 
posterior gonal process prominent with api- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Scanning electron micrographs of Lemnaphila scotlandae. 11, Egg, dorsal view. 12, Same, 


cal % parallel-sided, apex rounded, ventral- 
ly projected setula about *% distance from 
connection with subepandrial plate; anterior 
gonal process in lateral view (Fig. 8) a 
small, narrow, tapered process about % 
length of surstylus, bearing apical setulae; 
hypandrium (Fig. 10) narrowly V-shaped. 
Type Material.—The holotype male is la- 
beled ‘‘d/Fall Creek, Ithaca, N[ew]. 
Y[ork]./M. B. Scotland VII 25—46 [sic; Jul 
25—26],1933/TYPE No. 6501 Lemnaphila 
SCOTLANDAE E.T. Cresson, Jr, [red; 


VOLUME 102, NUMBER 3 


number and species name handwritten].”’ 
The holotype is double mounted (glued to 
a pin in a rectangular block of cork), is in 
excellent condition (slightly teneral), and is 
deposited in the ANSP (6501). 

Other Specimens Examined.—NEARC- 
TIC. CANADA. ONTARIO. Guelph, 12 Jan 
1976, J. M. Cumming (14 6, 16 2; GUE); 
16 Jan 1976, J. M. Cumming (5 2°; GUE); 
8 Aug 1978, W. A. Attwater (3 36,5 2; 
GUE); Limehouse, 1 Sep 1974, R. E. 
Roughley (1 3, 1 2; GUE); 2 Sep 1974, 
RES Roughley (1 6,5. 2: CNC, GUE); 6 
Oct 1974, R. E. Roughley (on Lemna minor 
in Black Creek), (1 2; GUE); Ottawa, 1967, 
J. G. Chillcott (2 36, 2 2; CNC); Wainfleet 
Bog (8 km S Welland), 14—28 Sep 1987, 
A. Stirling (2 6; WSU). 

UNITED STATES. FLORIDA. Alachua: 
Gainesville, 13—31 Oct 1986 (Lemna val- 
diviana: pools at Division of Plant Indus- 
try), G. R. Buckingham (4 6, 14 &, 17; 
JEPC); Pierce’s Homestead (Malaise trap; 
S9-TI1OS-RI8E), 11 May 1974, W. H. 
Pierce (1 2; USNM). Broward: Ft. Lauder- 
sale (Lemna valdiviana), 14 Nov 1988, G. 
R. Buckingham (1 6; USNM). Putnam: 
Rodman Reservoir (Lemna minor), 6 Oct 
1987, G. R. Buckingham (1 6, 1 Q&; 
USNM). Sumter: Lake Okahumpka (Lemna 
valdiviana), 22 Oct 1987, G. R. Bucking- 
ham (1 2; USNM). MICHIGAN. Clinton: 
Bath, 10 Oct 1965, (1 2; USNM). NEW 
YORK. Tompkins: Ithaca, Fall Creek, 25— 
26 Jul 1933, M. B. Scotland (12 6, 11 &; 
allotype; AMNH, ANSP, BMNH, CU, 
USNM); Ithaca, Sep 1933, M. B. Scotland 
(7 6, 11 2; CU, USNM). OHIO. Portage: 
Kent (near Kent State University stadium), 
23 May—4 Aug 1986, 1987, J. Edmiston 
Mi2ad, 12625 JEPC):Kent.24 Jun=2°Sep 
1987 (lab reared), J. Edmiston (9 3, 14 @; 
JEPC). Franklin: Columbus (Mirror Lake), 
23 Aug 1919, W. C. Kraatz (1 6; OHSU). 
Geauga: LaDue Reservoir (41°22.2'N, 
80°13.4'W; duckweed), 12 Sep 1976, B. A. 
Steinly (32 36, 21 2; USNM). 

Distribution.—WNearctic: Canada (ON), 
USA (FL, IL, MI, NY, OH). 


667 


Natural History.—Adult L. scotlandae 
are commonly found on or flying over the 
surface of floating duckweed thalli. The 
adults utilize the Lemna for food, leaving 
characteristic elongate gouges on the sur- 
face of the thallus. Seasonal distribution 
and overwintering stages are unknown; 
adults, however, have been collected in late 
November, suggesting that adults are able 
to survive the cold season like other north- 
temperate Ephydridae. Adults collected 
from the field and reared in the laboratory 
lived between 2—21 days (n = 13). 

Females lay eggs on the margin of a 
Lemna thallus, usually one egg per thallus. 
Females collected from the field and reared 
in the laboratory laid up to 41 eggs per day, 
averaging 21—26 eggs per day (n = 11) dur- 
ing their first four days in the laboratory. 
One female laid 334 eggs during 20 days. 

Eggs hatch in 4—6 days (x = 4.21; n = 
10), after which the first-instar larvae begin 
feeding on thallus tissue. Burrowing 
through the duckweed, the first-instar lar- 
vae feed for 1-3 days (x = 1.92; n = 10) 
before molting; second-instar larvae feed 
for 1-4 days (x = 2.62; n = 10) before 
molting; third-instar larvae feed for 2—7 
days (x = 4.85; n = 10) before pupariation 
occurs within the hollowed-out duckweed 
thallus. The pupal period ranged between 
6-8 days (x = 7.15 days; n = 10). 

Although the immatures and adults of 
this species are very tiny, three hymenop- 
terous parasatoids, as noted in the synony- 
my, have been reared from puparia (Mue- 
sebeck 1939, Scotland 1939, Buckingham 
1987). 


DESCRIPTION OF IMMATURE STAGES 


Egg (Figs. 11—12).—Length 0.35-0.37 
mm (x = 0.36; n = 12); width 0.13—0.17 
(x = 0.16; n = 12); ovoid, ends bluntly 
rounded; micropylar end porous; 4 longi- 
tudinal ridges dorsally with parallel rows of 
minute pores between ridges; flattened ven- 
trally with medioventral rows of minute 
pores and laterally with parallel rows of mi- 
nute pores. Chorion light brown. 


668 


Mature third-instar larva (Figs. 20- 
21).—Length 1.11—1.34 (x = 1.24; n = 5); 
width 0.22—0.32 (x = 0.28; n = 5). ver- 
miform, 12 segmented, white. Anterior end 
blunt with ventrally projected mandibles, 
posterior end with medioventrally projected 
spiracles. Integument covered with minute 
setulae. Cephalic segment (segment 1, 
pseudocephalic segment): antenna 2 seg- 
mented with rounded socket, proximal an- 
tennal segment minute, circular; distal an- 
tennal segment ovoid, greater than 10X 
length of proximal segment, lateroventrally 
with rounded sensory pits. 

Segment 2 (prothoracic segment): ven- 
trally and laterally with rows of comblike 
setae, projected dorsomedially over cephal- 
ic segment. Segment 3: longest segment, 1 
¥% length of segment 2 with rows of comb- 
like setae. Segments 3—11 similar, rounded, 
ventral surface with creeping welts; welts 
of segments 3 and 4 small; welts of seg- 
ments 5—11 large. Segment 12 with perianal 
pad ventrally and posterior spiracles ventro- 
medially; perianal pad bilobed, each lobe 
ovoid; posterior spiracle with cylindrical 
base and conical projection; conical projec- 
tion approximately twice as long as cylin- 
drical base and tapered to point. 

Cephalopharyngeal skeleton (Figs. 17— 
19): A-PDC 0.20-0.21 mm (n = 7), A- 
VDC 0.18—-0.20 mm (n = 7), PDC-VDC 
0.08 mm (n = 7). Mandibles fused dorsally, 
anteriorly pointed, posteriorly broadened. 
Hypopharyngeal sclerite minute, posterior- 
ly rounded, anteriorly slightly indented. 
Tentoropharyngeal_ sclerite anteriorly 
roundly pointed with anteroventral projec- 
tion, dorsal cornu posteriorly sharply point- 
ed, ventral cornu posteriorly broadened. 
First- and second-instar larvae similar. Dis- 
tinguishable by size of tentoropharyngeal 
sclerite. 

Puparium (Fig. 21).—Length 1.17—1.36 
(x = 1.28; n = 13); width 0.53-—0.60 (x = 
0.56; n = 13); width broadest posterome- 
dially in ventral view, tapering at each end; 
anterior end slightly curved ventrally, 
broadly rounded with remnants of cephal- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


opharyngeal skeleton clearly visible; pos- 
terior end slightly curved ventrally, spiracle 
remnants dark, perianal pad remnants visi- 
ble; margins of puparium smooth. Color 
uniformly pale brown except for black ce- 
phalopharyngeal skeleton remnants. 
Remarks.—This species is commonly 
called the Lemna fly (Scotland 1934, 1939, 
1940) or the duckweed miner, and the 
adults and immature stages live entirely in 
and around duckweed (Lemnaceae). Miss 
Minnie B. Scotland (1933), after whom the 
species was named, first reared L. scotlan- 
dae from duckweed (Lemna minor L.) that 
was collected in a fish hatchery at Fall 
Creek near Ithaca, New York. The species 
is now known to occur from southern Can- 
ada to northern Florida and west to Illinois. 


Lemnaphila wirthi Lizarralde de Grosso 
(Figs. 22—26) 


Lemnaphila wirthi Lizarralde de Grosso, 
1977:163; 1978:18-19 [review, descrip- 
tion of male terminalia]; 1989:55 [re- 
view, Argentina fauna].—Mathis and Za- 
twarnicki, 1995:97 [world catalog]. 


Diagnosis.—This species is distinguished 
from congeners by the following combina- 
tion of characters: Antenna entirely dark 
brown; length of Ist flagellomere only 
slightly longer than height; face in lateral 
view shallowly concave; ventral third of 
face densely microtomentose, blackish 
brown, concolorous with dorsal portion of 
face; scutellum densely microtomentose, 
appearing velvety, dark brown; katepister- 
num brown; coxae, femora, and tibiae yel- 
low. 

Description.—Minute to very small 
shore flies, length 0.95—1.20 mm. 

Head: Frons with mesofrons, parafrons, 
and anterior half of fronto-orbit concolo- 
rous, densely microtomentose, brownish 
black, appearing velvety; posterior fronto- 
orbit (from base of outer vertical seta to 
base of lateroclinate fronto-orbital seta) 
with whitish gray microtomentum. One 
fronto-orbital seta well developed, length 


VOLUME 102, NUMBER 3 669 


19 


20 


Figs. 17-21. Lemnaphila scotlandae. 17, Cephalopharyngeal skeleton, first-instar larva. 18, Same, second- 
instar larve. 19, Same, third-instar larva. 20, Third-instar larva, lateral view. 21, Puparium, ventral view. 


subequal to outer vertical seta, orientation like; length of outer vertical seta about 7 
postero-obliquely lateroclinate to laterocli- that of inner vertical seta, outer vertical seta 
nate; other fronto-orbital setae anterior of distinctly curved laterally, inner vertical 
large seta, at best weakly developed, hair- seta very shallowly curved, nearly vertical; 


670 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 22-26. Lemnaphila wirthi. 22, Male terminalia (epandrium, cercus, surstylus, aedeagus, aedeagal apo- 
deme), posterior view. 23, Same (with addition of postgonite), lateral view. 24, Internal male terminalia, lateral 
view. 25, Postgonite and subepandrial plate, ventral lateral view. 26, Fifth sternite and hypandrium, ventral view. 


ocelli arranged in isosceles triangle, with 
distance between posterior ocelli greater 
than between either posterior ocellus and 
anterior ocellus. Antenna entirely dark 


brown; length of Ist flagellomere only 
slightly longer than height; arista with 6-7 
long, dorsal rays. Face with shallowly con- 
cave in lateral view; ventral third of face 


VOLUME 102, NUMBER 3 


densely microtomentose, concolorous with 
frons; dorsal portion of face with very shal- 
low carina, mostly black except shiny black 
carina. Eye ratio 0.68—0.74. Gena-to-eye ra- 
tio 0.09—0.11. Maxillary palpus blackish 
brown; mouthparts generally brownish 
black. 

Thorax: Mostly brownish black to black; 
scutum subshiny, sparsely microtomentose; 
scutellum with dense microtomentum, 
brownish black, not shiny, but velvety ap- 
pearing like frons; pleuron mostly brownish 
black to black except dorsal half of anepis- 
ternum from some angles appearing silvery 
gray. Chaetotaxy: anterior dorsocentral seta 
inserted behind level of transverse suture; 
acrostichal setulae sparse anteriorly, in 2 
rows, lacking posteriorly; basal scutellar 
seta about % length of apical seta; katepis- 
ternal seta well developed, length subequal 
to posterior anepisternal seta. Wing: length 
1.08—1.24 mm; width 0.45—0.57 mm; vein 
R,,; mostly straight to very shallowly 
bowed anteriorly; costal vein ratio 1.5— 
1.73; length of costal section II about half 
of section III; M vein ratio 0.44-0.49. Hal- 
ter knob bright yellow; stem brownish yel- 
low. Legs: yellow except for brown 5th tar- 
someres. 

Abdomen: Dark brown, male 5th tergite 
subshiny; 5th sternite of male (Fig. 26) 
broadly and shallowly U-shaped, thickly 
formed, bearing numerous setulae along 
posterior margin and on membrane imme- 
diately adjacent. Male terminalia (Figs. 22— 
24): Epandrium (Figs. 22—23) comparative- 
ly well developed as a phalange at ventro- 
basal angle of cercus; cerci very well de- 
veloped, in lateral view (Fig. 23) irregularly 
hemispherical around posteromedial mar- 
gin, basolateral margin even, nearly straight 
in lateral view, shallowly concave in pos- 
terior view; surstylus (Figs. 22—23) long, 
slender, mostly parallel sided, apical % well 
sclerotized, bearing several setulae along 
length, apex rounded; subepandrial plate 
(Figs. 24—25) evenly arched, slender, band- 
like in lateral view (Fig. 24); aedeagus an- 
gulate in lateral view (Fig. 24), basiphallus 


671 


in lateral view somewhat quadrate with dis- 
tiphallus extended from ventral portion of 
basiphallus, distiphallus long, slender, shal- 
lowly sinuous, apex recurved; aedeagal 
apodeme (Fig. 24) enlarged, in lateral view 
semioval with keel prominent, wide; post- 
gonite in lateral view long and conspicu- 
ously sinuous, base moderately slender, api- 
cal % more slender, apex acutely pointed, 
apices cruciate in ventral view (Fig. 25); 
hypandrium in ventral view (Fig. 26) nar- 
rowly V-shaped with arms slightly flared 
apically, base wide and bluntly rounded. 

Type Material.—The holotype female is 
labeled ““ECUADOR. Manabi[:] Estero 
Balsa 9 Sept 1955/Collr.Levi-Castillo/Lem- 
naphila 2 n sp. det WWirth °60 [?]/LEM- 
NAPHILA WIRTHI L. DE GROSSO 
[handwritten]./HOLOTYPO [red].’’ The 
holotype is double mounted (glued to a pa- 
per triangle), is in good condition (right 
wing in glue, some setae broken), and is 
deposited in the USNM. 

Other Specimens Examined.—NEO- 
TROPICAL. DOMINICAN REPUBLIC. La 
Vega: Jarabacoa (1-2 km S; 19°06.9'N, 
70°37'W; 520 m), 8-21 May 1995, 1998, 
W. N. Mathis (296, 132; USNM). 

Distribution.—Neotropical: Argentina 
(Tucuman), Ecuador, West Indies (Domini- 
can Republic). 

Remarks.—The specimens from the Do- 
minican Republic are a major extension in 
the known range of this species, and em- 
phasizes the apparent lack of collecting this 
species and Lemnaphila generally. We now 
know this species from Argentina in the 
south to Ecuador and the Dominican Re- 
public in the north. 


PHYLOGENETIC CONSIDERATIONS 


Lemnaphila, as discussed previously (ge- 
neric diagnosis and discussion), is similar 
and related to Hydrellia and Cavatorella, 
and these three genera comprise the tribe 
Hydrelliini in the subfamily Hydrelliinae. 
The tribe Hydrelliini is distinguished from 
other tribes of Hydrelliinae by the follow- 
ing characters: specimens usually dull and 


672 


densely microtomentose (some species sec- 
ondarily sparsely microtomentose, subshi- 
ny, blackish); ocellar setae seldom as strong 
as pseudopostocellar setae, usually much 
weaker; eye bearing numerous short, inter- 
facetal, fluted setulae; postsutural supra-alar 
seta usually short, not longer than posterior 
notopleural seta; posterior notopleural seta 
at same level as anterior seta (independently 
and secondarily inserted above level of an- 
terior seta in Lemnaphila, as in Atissini); 
costa extended to vein M; midtibiae lacking 
dorsal, spinelike setae. 

Within Hydrelliini, the three included 
genera are distinguished from each other by 
the characters noted in the generic key (p. 
654). The relationships among these genera, 
however, are unresolved and remain prob- 
lematic. For example, although Lemnaphila 
and its included species are a well-corrob- 
orated, monophyletic lineage (see below), 
this lineage may eventually be found to be 
an included lineage within Hydrellia, i.e., 
Hydrellia lacks characterization by syna- 
pomorphies and its monophyly is unsub- 
stantiated. Cavatorella may likekwise be an 
included lineage within Hydrellia, perhaps 
closely related to Lemnaphila. 

Synapomorphies that conclusively estab- 
lish the monophyly of Lemnaphila are: 1. 
Ocellar setae lacking (also lacking in some 
species of Hydrellia); 2. Prescutellar acros- 
tichal seta lacking; 3. Vein R,,, short with 
costal section II shorter than section III (a 
shortened R,,, also occurs in some species 
of Hydrellia and in Cavatorella); 4. Poste- 
rior notopleural seta inserted near dorsal (su- 
tural) angle, clearly higher than the level of 
the anterior seta; 5. Anepisternum bearing a 
single seta along the posterior margin; 6. 
Epandrium weakly developed, incomplete 
dorsally, with membranous gap between lat- 
eral extensions, these extensions are as nar- 
row, lateral phalanges at base of cerci; 7. 
Cerci fused with medial margin of epan- 
drium; 8. Surstylus reduced to a simple, 
small process at the ventral margin of the 
fused epandrium/cerci complex; 9. Larvae 
mining the thalli of Lemnaceae (Hydrellia 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


albilabris (Meigen) from the Old World also 
mines the thalli of Lemna (Mathis and Za- 
twarnicki 1995)). The mining habit of H. al- 
bilabris in Lemna apparently arose indepen- 
dently. We have examined specimens of H. 
albilabris to verify that they are not closely 
related to Lemnaphila. 

In the presentation on species-level rela- 
tionships that follows, the characters used 
in the analysis are noted first. Each char- 
acter is immediately followed by a discus- 
sion to explain its states and to provide per- 
spective and any qualifying comments 
about that character. After presentation of 
the information on character evidence, an 
hypothesis of the cladistic relationships is 
presented and briefly discussed. The clad- 
ogram (Fig. 27) is the primary mode to con- 
vey relationships, and the discussion is to 
supplement the cladogram and is intended 
only to complement the latter. In the dis- 
cussion of character data, a “0” indicates 
the state of the outgroup; a “1” or “2” in- 
dicate the respectively more derived states. 
All multistate characters (1, 4, 8, 11, 14, 18, 
20) were treated as nonadditive characters 
(—), and characters 5, 6, and 9, which are 
autapomorphies for L. longicera, were 
made inactive (]) for the analysis, such as 
calculation of the consistency index. The 
numbers used in the presentation are the 
same as those on the cladogram (Fig. 27), 
and the sequence is the same as noted in 
the character matrix (Table 1). 

The other two genera of Hydrelliini, Hy- 
drellia and Cavatorella, were selected as 
outgroups. As the exemplar of Hydrellia, 
we chose the widespread type species of the 
genus, H. griseola (Fallén), and for Cava- 
torella, which is monotypic, we studied C. 
spirodelae Deonier, which occurs in China 
and Japan (Deonier 1995). Hydrellia is rep- 
resented by over 200 species worldwide 
(Mathis and Zatwarnicki 1995). 


CHARACTERS USED IN THE PHYLOGENETIC 
ANALYSIS 


Head: 


1. Vestiture of mesofrons: moderately mi- 


VOLUME 102, NUMBER 3 


Figs. 27. 


Hydrellia griseola 


Cavatorella spirodelae 


11 13 16 17 21 22 23 24 


crotomentose, appearing mostly dull 
(0); sparsely microtomentose, subshiny 
(1); densely microtomentose, appearing 
velvety black or whitish (2); meso- 
frons, except ocellar triangle, bare, 
shiny (3). This multistate character is 
treated as nonadditive. 


. Ocellar setae: present (laterad of an- 


terior ocellus, smaller than pseudopos- 
tocellar setae) (0); lacking or greatly re- 
duced (1). 


. Arrangement of ocelli: ocelli forming 


an isosceles triangle with distance be- 


Table 1. Matrix of characters and taxa used in the 


cladistic analysis of Lemnaphila (numbers for charac- 
ters correspond with those used in the text). 


OOOO0O0O00O 11111111112 222 2 

123456789 01234567890 123 4 
Hydrellia OOOO00000 OOODODOOOO000 OOO O 
Cavatorella 000100000 10000000000 O00 O 
L. longicera 111311001 02112011210 111 ? 
L. grossoae 210100110 11010121001 111 1 
L. scotlandae 210200100 02010111102 111 1 
L. wirthi 210100120 11011101000 111 1 
L. neotropica 210100110 11010101001 111 1 
L. lilloana 310100110 11010101001 111 1 


SIA 5. (6) 9) Zea 4 ato 


10 11 16 


673 


Lemnaphila longicera 


Lemnaphila wirthi 


16 
Lemnaphila grossoae 


Lemnaphila neotropica 


Lemnaphila lilloana 


Cladogram depicting hypothetical cladistic relationships among species of Lemnaphila. 


tween posterior ocelli about twice that 
between either posterior ocellus and 
anterior ocellus (0); ocelli forming an 
equilateral triangle or with distance be- 
tween posterior ocelli only slightly 
more than between either posterior 
ocellus and anterior ocellus (1). 


. Fronto-orbital setae: anterior fronto- 


orbital seta proclinate, smaller than 
posterior seta, which is laterobliquely 
reclinate (0); posterior seta elongate, 
oriented laterally (1); both setae re- 
duced but evident (2); both setae great- 
ly reduced or lacking (3). This multi- 
state character is treated as nonadditive. 


. Color of antenna: generally unicolo- 


rous, most brownish black but some- 
times yellowish (0); scape, pedicel, and 
basal half of Ist flagellomere yellow 
(1). The derived state of this character 
is an autapomorphy for L. longicera 
and is therefore treated as an inactive 
character in the analysis. 


6. Length of Ist flagellomere: about equal 


to or slightly greater than height (0); 
twice height (1). The derived state of 
this character is an autapomorphy for 


674 


Lh 


. Shape of face in lateral view: 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


L. longicera and is therefore treated as 
an inactive character in the analysis. 
face 
straight (0); shallowly concave, usually 
with a shallow, transverse groove (1). 


. Facial vestiture: face mostly uniform- 


ly microtomentose, color varying, de- 
pending on angle of view, except for 
narrow, vertical stripe (0); ventral third 
of face densely microtomentose, sil- 
very white, distinctly contrasted with 
blackish brown dorsal portion (1); ven- 
tral third of face densely microtomen- 
tose, dark brown (2). This multistate 
character is treated as nonadditive. 
Length of outer vertical seta: about *4— 
¥% length of inner vertical seta (0); less 
than half inner vertical seta (1). The de- 
rived state of this character is an auta- 
pomorphy for L. longicera and is there- 
fore treated as an inactive character in 
the analysis. 


Thorax: 


10. 


UY. 


|p 


PS: 


14. 


Position of anterior dorsocentral seta: 
inserted posterior of transverse suture 
(QO); inserted at level of transverse su- 
ture (1). 

Acrostichal setulae: numerous setulae 
and 1 well-developed pair of prescutel- 
lar setae (0); sparse setulae anteriorly, 
in 2 rows, lacking posteriorly, includ- 
ing prescutellar pair (1); lacking (2). 
This multistate character is treated as 
nonadditive. 

Comparative length of basal scutellar 
seta: long, from % to being subequal 
to length of apical seta (0); short, about 
¥% length of apical seta (1). 

Position of posterior notopleural seta: 
inserted at about same level as anterior 
seta (O); inserted at elevated position 
relative to anterior seta (1). 

Scutellar vestiture: moderately densely 
microtomentose, not appearing velvety 
(O); scutellum densely microtomentose, 
somewhat appearing velvety (1); scu- 
tellum densely microtomentose, ap- 
pearing velvety, dark brown, similar to 


LS! 


16. 


ie 


18. 


Nee 


20. 


notopleural patch (2). This multistate 
character is treated as nonadditive. 
Length of costal sections IIT and III: 
costal sections II and III about equal in 
length, vein R,,, relatively long (0); 
costal section III considerably longer 
than II, vein R,,, relatively short (1). 
Shape of anal angle of wing: deeply 
rounded, greatest depth equal to or 
greater than distance between apices of 
vein R,,; and M (0); very shallow 
rounded, greatest depth conspicuously 
less than distance between apices of 
vein R,,; and M (1). 

Number of setae and setulae along pos- 
terior margin of anepisternum: | large 
seta and 1-2 setulae (0); 1 seta, setulae 
greatly reduced or lacking (1). 
Presence or absence of katepisternal 
seta: seta well developed, length sub- 
equal to posterior anepisternal seta (0); 
seta greatly reduced (1); or lacking (2). 
This multistate character is treated as 
nonadditive. 

Color of katepisternum: brown (0); 
whitish yellow, concolorous with fore- 
coxa (1). 

Color of legs: legs entirely yellowish, 
with coxae whitish yellow and apical 
half of tibiae slightly brownish yellow 
(0); femora brown, tibiae and most tar- 
someres yellow (1); legs unicolorous, 
brown (2). This multistate character is 
treated as nonadditive. 


Abdomen: 


Ze 


Pio 


25: 


Development of epandrium at dorsum: 
dorsum of epandrium well developed 
to weakly developed (0); epandrium in- 
complete dorsally with membranous 
gap between lateral phalanges (1). 
Condition of cercus relative to epan- 
drium: cerci free from epandrium (0); 
cerci fused laterally to medial margin 
of epandrial phalange (1). 
Development of surstyli: generally well 
developed structures at ventral margin 
of epandrium (0); reduced to simple 


VOLUME 102, NUMBER 3 


675 
Table 2. Analysis of characters based on the cladogram (Fig. 27). 
Character 1 2 8 4 5 6 7 8 9 10 11 12 
Steps 3 1 i ) 1 1 1 2 1 1 2 1 
Con. Index 100 100 100 100 100 100 100 100 100 100 100 100 
Ret. Index 100 100 100 100 100 100 100 100 100 100 100 100 
Character 13 14 15 16 17 18 19 20 21 22 23, 24 
Steps 1 2 1 4 1 2 1 2 1 1 1 1 
Con. Index 100 100 100 50 100: 100 100 100 100 100 100 100 
Ret. Index 100 100 100 0) 100 100 100 100 100 100 100 100 


small process at ventral margin of the 
fused epandrial/cercal complex (1). 


Behavior: 


24. Feeding preference: miner in various, 
mostly aquatic plants (0); miner in the 
thalli of Lemnaceae (1). Lacking rear- 
ing records for L. lonicera, we coded 
this character as a ? for this species. 
Like all other members of Lemnaphila, 
however, we anticipate and predict that 
this species will also be found to be a 
miner in aquatic plants of Lemnaceae. 


ANALYSIS 


Using the implicit enumeration (ie*) op- 
tion of Hennig86, four most and equally 
parsimonious trees were generated, each 
with a length of 34 steps and consistency 
and retention indices of 0.91 and 0.86 re- 
spectively. The basal nodes in each of these 
cladograms are identical, with variation 
only in the more derived four species (the 
apical 2—3 nodes). 

The matrix was then subjected iteratively 
to successive weighing (xs w, ie*, cc) to 
determine a character’s contribution or 
weight and to find cladograms supported by 
the most consistent characters (Carpenter 
1988, Dietrich and McKamey 1995). Suc- 


cessive weighing produced a single clado- 
gram (Fig. 27) that is identical to one of the 
first four and is our cladogram of choice. 
The analysis of the characters for this clad- 
ogram is given in Table 2 and the weights 
of the various characters is given in Table 3. 

Hydrellia was consistently the basal out- 
group, with Cavatorella as the more im- 
mediate outgroup to Lemnaphila. Although 
the placement of Cavatorella as the sister 
group to Lemnaphila is currently the best 
indication of cladistic relationship, we do 
not attribute great weight to this relation- 
ship largely because only two synapomor- 
phies (characters 2 and 4) were found to 
support this sister-group relationship. 

The two basal lineages within Lemna- 
Phila are first L. longicera, which is the sis- 
ter lineage to the remaining species in the 
genus, followed by L. scotlandae. The next 
four species, which are all Neotropical in 
distribution (often at the same habitat), are 
obviously closely related and are very sim- 
ilar externally, and in the four equally par- 
simonious trees, these four species varied 
in position with respect to each other. In 
two of these trees, L. wirthi was basal to 
the other three, and in the other two trees, 
L. grossoae was the basal lineage. The sin- 
gle tree from successive weighing, howev- 


Table 3. Status (i.e., nonadditive —; inactive ]) and weights of characters after successive weighing. 


Character No. 1 2 3 4 5 
Weight, status IO= | WOE IE TOs | IG |[  IKORS | 
Character No. 13 14 NS) 16 iL7/ 
Weight, status Oar | WO= |[ MORE I Oke |p Ose I 


6 7 8 9 10 11 12 
OSS) TOS = 7 eee) See SP aoe 
18 19 20 21 oy) 23 24 
10 op Oe | Os 10s pO Ge | 


676 


er, has L. wirthi as the basal lineage with 
the other three species forming an unre- 
solved trichotomy (Fig. 27). 


ACKNOWLEDGMENTS 


We gratefully acknowledge the assistance 
and cooperation of many organizations and 
individuals who contributed to the field 
work and production of this paper. To Dr. 
David A. Grimaldi (AMNH), Mr. Donald E 
Azuma (ANSP), Mr. John E. Chainey 
(BMNH), Dr. James K. Liebherr (CU), Dr. 
Steve Marshall (GUE), Dr. Mercedes Li- 
zatralde de Grosso (IML), Dr. Norman 
Johnson (CHSU), Dr. Gustavo Spinelli 
(UNLP), Dr. Richard S. Zack (WSU) and 
their institutions, who loaned specimens, 
we express our sincere thanks. 

The illustrations were carefully inked by 
Mr. Young T. Sohn. Dr. Alan K. Graham, 
Kent State University, assisted with the 
preparation of the scanning electron micro- 
graphs (SEMs), and Mr. George L. Venable 
expertly produced the plate of SEMs (Figs. 
11-15). Dr. Stephen D. Gaimari prepared 
the final version of the cladogram (Fig. 27). 
For reviewing a draft of this paper we thank 
Drs. Stephen D. Gaimari, Benjamin A. 
Foote, and Richard S. Zack. 

Field work on St. Vincent, St. Lucia, and 
Dominica was supported by a grant from 
the Research Opportunity Fund, adminis- 
tered by David Pawson and Stanwyn G. 
Shetler, former Deputy Directors (USNM). 
In 1995, 1996, and 1998, field work on the 
West Indies was funded in large measure by 
grants from the Biodiversity Program (Bi- 
ological Surveys and Inventories, BSI), Na- 
tional Museum of Natural History, Smith- 
sonian Institution (Dr. Lynne R. Parenti, 
chair). Field work on the West Indies was 
greatly expedited through the able and 
pleasant assistance of my wife, N. Dianne 
Mathis, Ms. Hollis B. Williams, Mr. Kelvin 
Guerrero, Dr. Daniel E. Pérez-Gelabert, and 
Dr. Oliver S. Flint, Jr. 


LITERATURE CITED 


Buckingham, G. R. 1989. Lemnaphila scotlandae 
(Diptera, Ephydridae) and three of its parasites 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


discovered in Florida. Florida Entomologist 72(1): 
219-221. 

Carpenter, J. M. 1988. Choosing among multiple 
equally parsimonious cladograms. Cladistics 4(3): 
291-296. 

Clausen, P. J. and E. EK Cook. 1971. A revision of the 
Nearctic species of the tribe Parydrini (Diptera: 
Ephydridae). Memoirs of the American Entomo- 
logical Society 27: 1—150. 

Courtney, G. W., R. W. Merritt, H. J. Teskey, and B. 
A. Foote. 1996. Chapter 22. Aquatic Diptera. Part 
One. Larvae of Aquatic Diptera. Pp. 484-514. In 
R. W. Merritt and K. W. Cummins, eds., An in- 
troduction to the Aquatic Insects of North Amer- 
ica. 3rd edition, 862 pp. Kendall/Hunt Publishing. 
Dubuque, Iowa. 

Cresson, E. T., Jr. 1933. A new genus and species of 
the dipterous family Ephydridae reared from duck 
weed. Entomological News 44(9): 229-331. 

. 1944. Synopses of North American Ephydri- 
dae (Diptera). Parts IA and II. Transactions of the 
American Entomological Society 70: 159-180. 

Dahl, R. G. 1959. Studies on Scandinavian Ephydridae 
(Diptera Brachycera). Opuscula Entomologica, 
Supplementum 15: 1—224. 

Deonier, D. L. 1995. Cavatorella spirodelae Deonier 
(Diptera: Ephydridae), a new genus and new spe- 
cies from Spirodela (giant duckweed) in China 
and Japan. Insecta Mundi 9(3—4): 177-184. 

Deonier, D. L. and J. T. Regensburg. 1978. New re- 
cords of Ohio shore flies (Diptera: Ephydridae). 
Ohio Journal of Science 78(3): 154—155. 

Dietrich, C. H. and S. H. McKamey. 1995. Two new 
Neotropical treehopper genera and investigation 
of the phylogeny of the subfamily Membracinae 
(Homoptera: Membracidae). Proceedings of the 
Entomological Society of Washington 97(1): 1— 
16. 

Ferrar, P. 1987. A guide to the breeding habits and 
immature stages of Diptera Cyclorrhapha (Part 1: 
text). Entomonograph 8, 478 pp. E. J. Brill/Scan- 
dinavian Science Press. Leiden, Copenhagen. 

Foote, B. A. 1995. Biology of shore flies. Annual Re- 
view of Entomology 40: 417—442. 

Grimaldi, G. A. 1987. Phylogenetics and taxonomy of 
Zygothrica (Diptera: Drosophilidae). Bulletin of 
the American Museum of Natural History 186: 
103-268. 

Hennig, W. 1943. Ubersicht iiber die bisher bekannten 
Metamorphosestadien der Ephydriden, mit Neu- 


beschreibungen nach dem Material der Deutschen 
Limnologischen Sundaexpedition. (Diptera: Ephy- 
dridae). Arbeiten tiber morphologische und tax- 
onomische Entomologie aus Berlin-Dahlem 10 
(2-3): 105-138. 

Johannsen, O. A. 1935. Aquatic Diptera. Part II. Or- 
thorrhapha-Brachycera and Cyclorrhapha. Cornell 


VOLUME 102, NUMBER 3 


University Agricultural Experiment Station. Mem- 
oir 177: 1-62. 

Lizarralde de Grosso, M. S. 1977. Lemnaphila Cres- 
son, nuevo género para la region neotropical, con 
la description de dos especies nuevas. Limnobios 
1 (5): 159-164. 

. 1978. Nuevos aportes al conocimiento del gé- 

nero Lemnaphila Cresson (Diptera-Ephydridae). 

Neotropica 24 (71): 13-20. 

. 1980. Estados preimaginales de Ephydridae 

(Diptera) Argentinos, con clave de larvas. Physis 

(Buenos Aires), section C 39(96): 55—60. 

. 1989. Ephydridae de la Republica Argentina 
(Insecta-Diptera). Serie Monografica y Didactica 
No. 3. Facultad de Ciencias Naturales e Instituto 
Miguel Lillo Universidad Nacional de Tucuman. 
93 pp. 

Mansor, M. and G. R. Buckingham. 1989. Laboratory 
host range studies with a leaf-mining duckweed 
shore fly. Aquatic Plant Management 27: 115— 
118. 

Marsh, P. M. 1979. Family Braconidae. Pp. 144—295. 
In K. V. Krombein, P. D. Hurd, Jr, D. R. Smith, 
and B. D. Burks, eds., Catalog of Hymenoptera in 
America North of Mexico. Vol. 1, 1198 pp. Sym- 
phyta and Apocrita (Parasitica). Smithsonian In- 
stitution Press, Washington, D.C. 

Mathis, Wayne N. 1986. Studies of Psilopinae (Dip- 
tera: Ephydridae), I: A Revision of the shore fly 
genus Placopsidella Kertész. Smithsonian Contri- 
butions to Zoology 430: iv + 30 pp. 

Mathis, Wayne N. and Tadeusz Zatwarnicki. 1990a. A 
revision of the western Palearctic species of Athy- 
roglossa (Diptera: Ephydridae). Transactions of 
the American Entomological Society 116(1): 103— 
11333). 

. 1990b. Taxonomic notes on Ephydridae (Dip- 

tera). Proceedings of the Biological Society of 

Washington 103 (4): 891—906. 

. 1995. A world catalog of the shore flies (Dip- 
tera: Ephydridae). Memoirs on Entomology, In- 
ternational, Associated Publishers, Gainesville, 
FL, 4: vi + 423 pp. 

McAlpine, J. EF 1981. Morphology and terminology- 
adults, pp. 9-63. In McAlpine, J. EF, et al., eds., 
Manual of Nearctic Diptera, Vol. 1. Agriculture 
Canada Monograph 27, Ottawa, vi + 674 pp. 

Merritt, R. W., D. W. Webb, and E. I. Schlinger. 1996. 


677 


Chapter 23. Aquatic Diptera. Part Two. Pupae and 
adults of Aquatic Diptera. Pp. 515-548. In R. W. 
Merritt and K. W. Cummins, eds., An introduction 
to the Aquatic Insects of North America. 3rd ed., 
862 pp. Kendall/Hunt Publishing, Dubuque, Iowa. 
Muesebeck, C. E W. 1939. Three new hymenop- 
terous parasites of the Lemna fly. Proceedings of 
the Entomological Society of Washington 41 (3): 
58-62. 

. 1979. Family Diapriidae. Pp. 1127—1150. In 
K. V. Krombein, P. D. Hurd, Jr., D. R. Smith, and 
B. D. Burks, eds., Catalog of Hymenoptera in 
America North of Mexico. Vol. 1, 1198 pp. Sym- 
phyta and Apocrita (Parasitica). Smithsonian In- 
stitution Press, Washington, D.C.. 

Scotland, M. B. 1934. The animals of the Lemna as- 
sociation. Ecology 15(3): 290-294. 

. 1939. The Lemna fly and some of its parasites. 

Annals of the Entomological Society of America 

32: 713-718. 

. 1940. Review and summary of studies of in- 
sects associated with Lemna minor. Journal of the 
New York Entomological Society 48(4): 319-332, 
4 plates. 

Steinly, B. A., E. Lisowski, and D. Webb. 1987. The 
distribution of shore flies (Diptera: Ephydridae) in 
Illinois. Entomological News 98(4): 165-170. 

Wirth, W. W. 1965. Family Ephydridae. Pp. 734-759. 
In A. Stone, et al., eds., A Catalog of the Diptera 
of America north of Mexico. United States De- 
partment of Agriculture, Agriculture Handbook, 
276: iv + 1696 pp. 

Wirth, W. W., W. N. Mathis, and J. R. Vockeroth. 1987. 
98. Ephydridae. Jn J. FE McAlpine, ed., Manual of 
Nearctic Diptera, Vol. 2: 1027-1047. Monograph 
28, 675-1332 + iv pp. Research Branch, Agri- 
culture Canada, Ottawa. 

Wirth, W. W. and A. Stone. 1956. Chapter 14. Aquatic 
Diptera. Pp. 372-482. In R. L. Usinger, ed., 
Aquatic insects of California. 508 + x pp. Uni- 
versity of California Press, Berkeley. 

Zatwarnicki, T. 1992. A new classification of Ephy- 
dridae based on phylogenetic reconstruction (Dip- 
tera: Cyclorrhapha). Genus 3(2): 65-119. 

. 1996. A new reconstruction of the origin of 

Eremoneuran hypopygium and its implications for 

classification (Insecta: Diptera). Genus 7(1): 103— 

17/3) 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 678-687 


ON THE VALIDITY OF HELICONIUS TRISTERO BROWER AND 
HELICONIUS MELPOMENE MOCOA BROWER, WITH NOTES ON SPECIES 
CONCEPTS IN HELICONIUS KLUK (LEPIDOPTERA: NYMPHALIDAE) 


ANDREW V. Z. BROWER 


Department of Entomology, Oregon State University, Corvallis, OR 97331-2907, 
U.S.A. (e-mail: browera@bcc.orst.edu) 


Abstract.—Lamas’ (1998) criticisms of Brower’s (1996a) taxonomy are shown to be 
based on misinterpretations of the evidence and of the rules of nomenclature. The names 
H. amaryllis amaryllis f. bellula Stichel and H. melpomene bellula Turner are unavailable. 
The name H. bellula Brower is available but invalid. The name H. melpomene mocoa 
Brower is the valid name for the subspecies of H. melpomene from the Putumayo region 
of southeastern Colombia. The Genotypic Cluster Species Concept is contrasted unfavor- 
ably to the Phylogenetic Species Concept with respect to the aims of systematics in 
general, and the resolution of relationships among geographically differentiated Heliconius 


taxa in particular. 


Key Words: 
scription, subspecies 


Recently, Lamas (1998) published a crit- 
ical discussion of two new names applied 
to Heliconius butterflies from the upper Rio 
Putumayo basin in southeastern Colombia 
(Brower 1996a). The taxa in question are 
Miillerian mimics: one (H. tristero) was de- 
scribed as a species in the H. cydno Dou- 
bleday clade, while the other was named as 
a subspecies, H. melpomene mocoa. Most 
members of the cydno clade exhibit blue- 
and-white or blue-and-yellow wing patterns 
and mimic species in the sara-sapho group 
(cf. Brown 1981, Brower 1994, and Brower 
and Egan 1997 for reviews of phylogenetic 
hypotheses among Heliconius species), so 
it was a surprise to discover a new species 
mimicking sympatric races of H. melpo- 
mene (L.) and H. erato (L.). 

That these formerly conflated taxa rep- 
resent two distinct entities is supported by 


Heliconius melpomene bellula, Heliconius tristero, nomenclature, circum- 


robust evidence stemming from three dif- 
ferent sources (mitochondrial DNA se- 
quences, genitalic morphology and wing 
patterns), and that at least one of them re- 
quired a new name is not at issue. Lamas’ 
criticisms of my paper focussed on the fol- 
lowing problems: (1) the availability of H. 
bellula Stichel (1923); (2) the supposed 
synonymy of H. melpomene mocoa Brower 
with bellula auctt.; (3) my interpretation of 
the International Code of Zoological No- 
menclature (1985) with regard to hybrids; 
and (4) the logic of my species concept. 
First, I will address these problems in nar- 
row relation to Lamas’ critique of my paper 
(1996a). My rebuttal of Lamas’ criticisms 
is followed by a brief discussion of the spe- 
cies problem in Heliconius. An English 
translation of the relevant section of Lamas’ 
paper is included as Appendix 1. 


VOLUME 102, NUMBER 3 


REPLY TO LAMAS’ CRITIQUE 


Is Heliconius bellula Stichel (1923) an 
available name? 


As Lamas (1998) pointed out, Stichel 
(1923) applied the name bellula to a form 
of H. amaryllis amaryllis C. & R. Felder 
(H. amaryllis is now considered to be a sub- 
species of H. melpomene; cf. Ackery and 
Smiles, 1976). Lamas argued that Stichel’s 
employment of bellula was infrasubspecific 
and therefore formally unavailable. How- 
ever, Neustetter (1929) cited Stichel’s name 
as a trinominal form of H. amaryllis, thus 
implying that the name was available 
(ICZN Article 16). The subsequent employ- 
ment of the name for a geographical race 
by more recent authors (e.g., Turner 1971, 
Brown 1979, Sheppard et al. 1985, Mallet 
1993, Brower 1996b) further suggests that 
the name has been treated as available (Art. 
45gii), and it was under that premise that I 
dealt with it in my paper (Brower 1996a)'. 

However, given that Stichel’s holotype is 
not a representative of the melpomene clade 
(see Brower 1996a and below), Lamas’ 
opinion that Stichel’s name was quadrinom- 
inal provides a convenient excuse to sink it 
and avoid the complications described 
above. If its original designation is deemed 
infrasubspecific, a name remains unavail- 
able until a description is provided (Art. 
10c). Lamas claimed that Turner (1971) 
employed the name bellula to refer to a 
subspecific entity, thereby becoming its au- 
thor (Arts. 23j and 50c). However, Turner’s 
tentative use of the name in a figure legend 
(followed by a question mark) was not ac- 
companied by a description. Likewise, 
Brown (1979), Sheppard et al. (1985), Mal- 


'T examined and discussed Stichel’s holotypes of 
bellula, permira and degener. Lamas suggested that I 
was unaware of four additional forms described by Sti- 
chel from the Mocoa region. Rather, these were delib- 
erately omitted from discussion, because it was clear 
from the original descriptions that they represent var- 
ious hybrid forms with recombinant wing patterns that 
are even less similar to the modern “‘bellula” concept 
than the two other forms I discussed. 


679 


let (1993) and Brower (1996b) used the 
name, but none of them provided a descrip- 
tion of the taxon, nor did any of them cite 
Stichel’s description (which would not per 
se change its availability, in any case: Art. 
11dii). All of those uses of bellula employ 
it as a nomen nudum and none of them sat- 
isfies the criteria of availability (Art. 13a). 

Holzinger and Holzinger (1994) provided 
a description and illustration of H. melpo- 
mene bellula and cited Stichel (1923), but 
their book is not consistently binominal (it 
uncritically employs numerous infrasubspe- 
cific names), and its contents are thus un- 
available (Art.11c). Ironically, it appears 
that my 1996 paper is the first to provide a 
description for H. bellula that would render 
it available under all the criteria of Art. 11 
(via illustrations, description of differenti- 
ating features, identification of a holotype, 
and citation of Stichel 1923). But because 
I circumscribed the concept narrowly, to re- 
fer to a hybrid form typified by Stichel’s 
original type specimen, the name is invalid. 
Even so, my definition prevents subsequent 
usage of bellula for any other Heliconius 
taxon (Art. 23h; see hybrid section, below). 
Thus, the correct authorship of Heliconius 
bellula is Brower (1996a), but the name 
does not refer to any valid taxon, regardless 
of its repeated misapplication in recent 
works (including my own). 


Subjective synonymy of bellula and 
mocoa? 


Lamas (1998) argued that H. melpomene 
mocoa Brower is a junior subjective syno- 
nym of H. melpomene bellula auctt. Since 
bellula is not a valid name, this is a moot 
point, but it raises another important issue: 
Lamas’ claim shows that has been deceived 
by Miillerian mimicry! It is the holotype of 
bellula and the holotype of H. tristero that 
are closely-related, while the holotype of H. 
melpomene mocoa represents a different 
clade. Had Lamas examined the relevant 
characters, he would have seen that bellula 
and mocoa are not the same under any rea- 
sonable circumscription H. bellula (what- 


680 


ever its status) is not an example of the H. 
melpomene species group; instead, it is 
close to H. tristero in the H. cydno species 
group. To include bellula and mocoa in the 
same species, one would have to view H. 
cydno Doubleday as a subjective junior 
synonym of H. melpomene (L.)! I described 
this problem explicitly in Brower (1996a), 
and it was precisely the desire to avoid such 
confusion that prompted me to coin new 
names for both the cydno relative (H. tris- 
tero) and the melpomene relative (H. mel- 
pomene mocoa) in the first place. 

Even if bellula and mocoa were closely 
related, it is clear from Stichel’s (1923) de- 
scriptions (Appendix 2) that he felt that bel- 
lula was different enough from the local 
“nominate form’ to warrant a separate 
name. Stichel believed (in error) that his 
three “‘typical’’ specimens were H. amaryl- 
lis amaryllis, today recognized as a distinct 
H. melpomene subspecies with an allopatric 
distribution in the Huallaga valley of Peru 
(Sheppard et al., 1985). Although ideas 
about what entities within Heliconius de- 
serve names have changed as knowledge of 
phenotypic variation and geographical dis- 
tribution has grown, efforts to redefine an 
old name of dubious availability in specific 
contradiction to its original author’s intent 
seem procrustean. A more reasonable view 
is that the “‘nominate form” of H. melpo- 
mene from Mocoa and environs was anon- 
ymous until I (Brower 1996a) named it. 


Hybrids and the ICZN 


Lamas (1998) also criticized as erroneous 
my invocation of Articles 1b and 23h to 
curtail the current usage of bellula because 
its holotype is a hybrid (Brower 1996a). He 
pointed out that because the definition of 
““‘hybrid”’ in the ICZN glossary states that 
offspring of crosses between conspecific 
subspecies are not hybrids, bellula cannot 
be considered a hybrid taxon. Re-exami- 
nation of Brower (1996a) shows Lamas’ 
point to be technically incorrect: I hypoth- 
esized that the bellula holotype specimen 
was the result of hybridization between H. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


heurippa and H. tristero, both of which I 
explicitly claimed were species in that pa- 
per. Under such circumstances ICZN article 
23h specifically applies’. 

Whether or not the bellula holotype ac- 
tually represents a hybrid or not (and which 
parental species might have spawned it) is 
difficult to determine in retrospect and 
without extensive data from experimental 
genetic crosses. That additional specimens 
displaying the same pattern have not been 
collected suggests that the holotype exhibits 
a rare recombinant wing pattern (typical of 
the phenotypic diversity found in other Hel- 
iconius hybrid zones). By contrast, there are 
multiple specimens from multiple localities 
that appear very similar to the holotype of 
H. melpomene mocoa, and illustrations of 
‘*H. melpomene bellula’’ in recent works 
(e.g., Turner 1971, Brown 1979, Holzinger 
and Holzinger 1994) all lack the yellow 
spots that are present in the bellula holo- 
type. My interpretation that the specimen 
does not represent a “pure’’ geographical 
race is complementary to Stichel’s (1923) 
opinion that the bellula holotype is transi- 
tional, as shown by his description of it as 
distinct and separate from his series of 
specimens of the local “‘nominate’’ form. 


Logical Consistency 

I made several statements that Lamas 
chose to overlook in his conclusion that my 
taxonomic argumentation (Brower 1996a) 
“‘no tiene sustento l6gico al ser refutable.” 
First, I did not argue that the current clas- 
sification of Heliconius is not illogical. In- 
deed, I stated that I chose the ranks I em- 
ployed to preserve nomenclatorial stability, 


*Lamas objected my use of the term “forbid” 
(translated as “‘prohibe”’ in his paper) with reference 
to the ICZN rule on names applied to interspecific hy- 
brids. Article 23h says, ““A species-group name estab- 
lished for an animal later found to be a hybrid . . . must 
not be used as a valid name for either of the parental 
species, even if it has priority over all other available 
names for them, but it may enter into homonymy.” I 
leave it to the reader to contemplate whether “must 
not be used”’ and “forbids the use of”? mean the same 
thing, or not. 


VOLUME 102, NUMBER 3 


even though I knew that the species delim- 
itations among geographically polymorphic 
Heliconius butterflies were effectively ar- 
bitrary. The final points of my paper were 
that, 


“The mitochondrial DNA data suggest 
that the degrees of relationship among the 
H. melpomene races and among the H. 
cydno races are similar, and that diver- 
gence times within each group are also 
approximately the same, implying that 
the taxonomic rank of each group should 
also be the same. The entire species-level 
classification of the genus will probably 
require revision as additional data be- 
come available.” 


Ignoring these qualifying remarks, Lamas 
offered (without providing new data or ex- 
amining the characters discussed in Brower 
[1996a]) a hypothesis of circumscription 
that he considered ‘“‘more valid’? than my 
view: that H. timareta Hewitson, H. heurip- 
pa Hewitson and H. tristero are conspecific. 
He based this notion on these entities’ geo- 
graphical proximity on the northeastern 
slope of the Andes, evidence of hybridiza- 
tion among them (which he denied on the 
following page), and their hypothesized 
close relationship (based on studies of 
mtDNA: Brower 1996a, 1996b). However, 
both the mtDNA studies and data from a 
more comprehensive recent analysis (Brow- 
er and Egan 1997) imply that H. timareta, 
H. heurippa and H. tristero are no more 
closely related to one another than any of 
them is to H. cydno or H. pachinus Salvin. 
Therefore, if one wanted to lump diagnos- 
ably different taxa based on the symple- 
siomorphy of potential interbreeding, the 
logically consistent choice would be to col- 
lapse all of these taxa under the oldest spe- 
cies-group name (H. cydno). I said as much 
in 1996a. 

If Lamas (1998) were as concerned with 
the consistent assignment of taxa to the ap- 
propriate rank as his criticism of Brower 
(1996a) implies, then according to his 
‘““more valid’? hypothesis, he should have 


681 


named his new Heliconius subspecies (de- 
scribed in the same paper) not “H. timareta 
timoratus, > but ““H. heurippa timoratus” 
(if timareta and heurippa are conspecific, 
then the former is either a junior synonym 
or a subspecies of the latter). Under the in- 
terpretation Lamas claimed that he prefers, 
timoratus would seem to be an infrasubspe- 
cific form of the subspecies H. heurippa ti- 
mareta! The complexity of sorting out 
names, ranks and relationships among geo- 
graphic races in Heliconius obviously pre- 
sents a challenge that neither Lamas nor I 
have yet resolved in a fully satisfactory 
manner. 


Synonymies 


Heliconius melpomene mocoa Brower, 
1996a. 

Heliconius melpomene bellula auctt. (Turn- 
er 1971, Brown 1979, Sheppard et al. 
1985, Mallet 1993, Brower 1994, Holzin- 
ger & Holzinger 1994) [misidentifica- 
tions; unavailable name]. 

Heliconius melpomene mocoa Brower 
1996a: 328. Holotype: Colombia, Dpto. 
Putumayo, 1—3 km N. Mocoa on rd. to 
Pitalito, 25 March 1992 leg. AVZ Brow- 
er. Deposited in Cornell University Insect 
Collection (examined). 


Heliconius heurippa X Heliconius tristero, 
natural hybrid 

Heliconius amaryllis amaryllis f. bellula 
Stichel, 1923: 262. Original type speci- 
men: Colombia, Rio Putumayo, Rio Gua- 
yuyaco, 7 July 1921 leg. W Hopp (see 
Brower 1996a for transcript of original 
labels). 

Heliconius amaryllis f. bellula Stichel; 
Neustetter 1929: 58. 

Heliconius amaryllis f. bellula Stichel; 
Brower 1996a [identified as hybrid H. 
heurippa X H. tristero]. 


NOTES ON SPECIES CONCEPTS IN HELICONIUS 


I suspect that the root of Lamas’ criti- 
cisms of my concepts of tristero and mocoa 
lies in his concept of species in Heliconius, 


682 


which differs from mine. The species prob- 
lem in Heliconius has been controversial 
for many years (e.g., Eltringham 1916 vs. 
Kaye 1916); indeed, it is the very complex- 
ity of geographical diversification within 
the genus that has led it to become a model 
system for the study of the evolutionary ge- 
netics of mimicry (Sheppard 1960, Emsley 
1964, Turner 1971, Brown et al. 1974, Mal- 
let 1993). Many of the Heliconius papers 
from the post-typological period applied the 
biological species concept (BSC; Mayr 
1940)), which unites allopatric taxa based 
on their potential to interbreed. In recent 
years, the BSC has been criticized on the- 
oretical and practical grounds, and numer- 
ous alternative species definitions have 
been proposed that offer more operational 
criteria for species delimitation (Eldredge 
and Cracraft 1980, Mishler and Donoghue 
1982, de Queiroz and Donoghue 1988, Nix- 
on and Wheeler 1990, Baum and Shaw 
1995, Mallet 1995). Lamas’ critique im- 
plied that he favors the “genotypic cluster” 
species concept (GCSC; Mallet 1995), 
while I (Brower 1996a) applied the phylo- 
genetic species concept (PSC) as elaborated 
by Nixon and Wheeler (1990). 


Evidence and Criteria of Specific 
Distinctness 


Mallet (1995) argued that the PSC fails 
to provide “‘clear guidelines” for dividing 
species because it recognizes groups based 
on apomorphy: ‘‘With detailed morphology 
and modern molecular techniques,”’ he as- 
serted, “‘one can find apomophies for al- 
most every individual,”’ which he suggested 
would result in rampant splitting and pro- 
liferation of species. This simplistic cari- 
cature of the cladistic method misrepresents 
the procedure of species delimitation, 
which has been explored in depth by Davis 
and Nixon (1992), Doyle (1995) and Brow- 
er (1999). Cladists identify populations of 
organisms that they hypothesize to be dis- 
tinct, and seek discrete differences between 
them. If they discover such differences, cla- 
dists consider the populations to be separate 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


species; otherwise, they are collapsed into 
a single species. That the cladistic approach 
may yield finer resolution of the hierarchi- 
cal pattern of diversity than alternative 
methods is considered by many to be an 
asset. In short, the guidelines of the PSC 
are clear: it is Mallet’s understanding of 
them that seems cloudy. 

Mallet’s (1995) alternative, the GCSC, 
views species as “‘identifiable genotypic 
clusters” recognizable by ‘a deficit of in- 
termediates”” at single and multiple loci, 
and views speciation as “‘the production of 
divergent populations that can coexist in 
sympatry.”’ There are a number of problems 
with this approach. First, there are no ex- 
plicit criteria for identification of genotypic 
clusters. That Mallet employed “deficit” 
instead of “‘absence’’ implies that he be- 
lieves that the clusters need not be fixed for 
alternate alleles, but merely differ in allele 
frequency by some “‘significant’’ amount. 
Further ambiguities include how many 
markers should be sampled (Mallet [1996] 
suggested multiple loci), and what should 
be done if one locus suggests continuity, 
while another suggests distinction. The 
across-locus averages that Mallet (1995) 
employed in the hybrid indices he presented 
would disguise heterogeneity among loci. A 
more sophisticated and logically consistent, 
but equally labor-intensive approach to 
multilocus species discrimination was pre- 
sented by Doyle (1995). In practice, how- 
ever, methods like these that rely on ex- 
haustive characterization of gene pools are 
rarely employed in the study of biological 
diversity, because they entail intensive, 
quantitative sampling that is not feasible in 
most circumstances. 

Another difficulty with the GCSC is non- 
dimensionality.—Maallet’s concept is only 
useful for contemporaneous taxa in sym- 
patry or parapatry. To cover everything 
else, Mallet (1995) suggested that “‘closely 
related allopatric forms should mostly be 
considered conspecific.’’ But how are we to 
determine that allopatric forms are “‘closely 
related?’’ If allopatric populations are 


VOLUME 102, NUMBER 3 


“identifiable,” then they logically satisfy 
Mallet’s GCSC criteria and are distinct spe- 
cies by definition; if they are not identifi- 
able, then why would we hypothesize that 
they were different in the first place? When 
the hypothetical taxa under investigation 
occur in allopatry, researchers are forced to 
turn to the empirical comparison of features 
of organisms to draw inferences about their 
taxonomic relationships. The PSC recogniz- 
es separate species only when fixed differ- 
ences are discovered between hypothesized 
groups, which will always result in the rec- 
ognition of a minimal number of taxa in 
comparison to methods based on frequency 
differences. The claim that the PSC over- 
splits taxa relative to the GCSC is simply 
false. 

In summary, Mallet’s GCSC is a meth- 
odologically explicit version of the nondi- 
mensional BSC (Mayr 1963). It is labor- 
intensive, depending upon sampling of mul- 
tiple individuals at multiple loci to provide 
empirical evidence of the absence of inter- 
breeding. It bases the decision of specific 
distinctness on an arbitrary and unstated 
level of phenetic bimodality in a histogram 
of average genetic scores. Although Mallet 
and colleagues have made a rather convinc- 
ing case for the distinctness of Heliconius 
erato cyrbia Godart and H. himera Hewit- 
son on the basis of this method (Jiggins et 
al. 1996), that work represents a laborious 
multi-year, multi-authored effort to corrob- 
orate a conclusion about a single pair of 
taxa that had already been hypothesized by 
systematists years before (Descimon and 
Mast-de Maeght 1984). How Mallet’s con- 
cept could be useful in the best of circum- 
stances to a museum taxonomist working 
with qualitative samples of preserved, dead 
specimens on pins is not clear. What is 
quite evident is that Lamas has never em- 
ployed the GCSC in any of his published 
taxonomic work, including the descriptions 
of new subspecies in Lamas (1998). Given 
this lack of consistency and rigor, it is es- 
pecially ironic that my (1996a) names 
should be subject to such scrutiny, when 


683 


they are perhaps the most thoroughly-di- 
agnosed Heliconius taxa that have been 
published, the differentiating characters 
having been drawn from the results of cla- 
distic analyses of mtDNA, and corroborated 
with diagnostic characters from external 
and internal morphology (Brower 1996a) 
and data from a nuclear gene (Brower and 
Egan 1997). 


Recognition and Circumscription of 
Subspecies 


Although the genus Heliconius contains 
numerous diagnosably different populations 
that bear valid species-group names, most 
of these are considered to refer to intraspe- 
cific variations. Since the rejection of ram- 
pant typological splitting in the early 20% 
Century (e.g., Riffarth 1902), Heliconius 
species have been circumscribed primarily 
by the BSC criterion of interbreeding. Un- 
der that criterion, otherwise uniform para- 
patric populations that hybridize where they 
abut have been considered conspecific. 
Likewise, distinct populations that hybrid- 
ize with each of two otherwise allopatric 
neighbors provide a transitive link that has 
allowed lumping of chains of populations 
into single, geographically extensive “‘bio- 
logical species.’ These species’ component 
*““geographical races’”’ are diagnosably dif- 
ferent (i.e., they display heritable characters 
that allow their unambiguous determina- 
tion), and have been considered by many 
researchers (e.g., Brown et al. 1974, Shep- 
pard et al. 1985, Brower 1996a, 1996b) to 
represent historically distinct entities. That 
their names are in common use in the lit- 
erature and in museum classification 
schemes is a de facto acknowledgement of 
their recognition as taxa, even by those bi- 
ologists who would emphatically deny their 
specific status. According to the PSC, diag- 
nosably distinct groups that it is useful to 
name are considered separate species, and 
every recognized “geographical race’? in 
Heliconius should be a phylogenetic spe- 
cies. 

The ICZN (Art. 45a) considers names at 


684 


both specific and subspecific levels to be 
labels for taxa of a single category, the Spe- 
cies Group. The concept of subspecies is 
simply a convenient label for taxa at one of 
the potentially many hierarchical levels 
nested within the genus, as revealed by cla- 
distic studies. The Code is sensibly silent 
on the problems of definition and boundary 
determination of concepts associated with 
name-bearing type specimens; such deci- 
sions are considered subjective, and left to 
the judgement of the describer and subse- 
quent employers of the name. In most in- 
stances, the older Heliconius names were 
described from one or a few dead speci- 
mens in European collections, and were not 
accompanied by any discussion of the cir- 
cumscription of the associated concept, be- 
yond designation of an intraspecific level 
(e.g., subspecies, form, aberration, etc.). A 
good example of such a perfunctory de- 
scription is Stichel’s original diagnosis 
(1923) of H. amaryllis amaryllis f. bellula 
(Appendix 2). 

Minimal original descriptions leave a 
great deal of latitude for subsequent inter- 
pretation. Such interpretations should strive 
to maintain nomenclatorial stability, but 
only when the names preserved are precise- 
ly and accurately associated with empiri- 
cally supported concepts. If a concept di- 
verges from the description due to the ac- 
quisition of new specimens and data to the 
point that the description is no longer ade- 
quate, some action is called for, ranging 
from redescription of the holotype to the 
separate description of differentiated con- 
cepts as distinct taxa. The currently-recog- 
nized “‘geographical races”’ (= phylogenet- 
ic species) of Heliconius are taxa corrobo- 
rated by a century of empirical research in 
laboratories, museums, and the field. They 
bear names originally applied to the single 
holotype or short type series, and to which 
the current concepts correspond. The focus 
of modern systematic effort should be on 
the empirical diagnosis of such taxa and the 
inference of hierarchical relationships 
among them, not on empty disputes over 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


the arbitrarily determined ranks that partic- 
ular taxa do or do not represent. H. mel- 
pomene mocoa and H. tristero are both 
diagnosably distinct taxa of the Species 
Group. 


ACKNOWLEDGMENTS 


Thanks to D. D. Judd and E C. Thomp- 
son for advice on the ICZN, A. Warren for 
checking my translation of Lamas’ Spanish, 
and A. Kretzer for help with Stichel’s Ger- 
man. Thanks to R. I. Vane-Wright for his 
thorough and candid reviews, and to Ger- 
ardo Lamas for making me play by the 
rules. 


LITERATURE CITED 


Ackery, P. R. and R. L. Smiles. 1976. An illustrated 
list of the type-specimens of the Heliconiinae 
(Lepidoptera: Nymphalidae) in the British Muse- 
um (Natural History). Bulletin of the British Mu- 
seum (Natural History) Entomology 32: 171-214. 

Baum, D. A. and M. J. Donoghue. 1995. Choosing 
among alternative “phylogenetic” species con- 
cepts. Systematic Botany 20: 560-573. 

Baum, D. A. and K. L. Shaw. 1995. Genealogical per- 
spectives on the species problem, pp. 289-303. In 
Hoch, P. C. and A. G. Stephenson, eds., Molecular 
and Experimental Approaches to Plant Biosyste- 
matics. Missouri Botanical Garden., St. Louis. 

Brower, A. V. Z. 1994. Phylogeny of Heliconius but- 
terflies inferred from mitochondrial DNA se- 
quences (Lepidoptera: Nymphalidae). Molecular 
Phylogenetics and Evolution 3: 159-174. 

. 1996a. A new mimetic species of Heliconius 

(Lepidoptera: Nymphalidae), from southeastern 

Colombia, as revealed by cladistic analysis of mi- 

tochondrial DNA sequences. Zoological Journal 

of the Linnean Society 116: 317-332. 

. 1996b. Parallel race formation and the evo- 

lution of mimicry in Heliconius butterflies: A phy- 

logenetic hypothesis from mitochondrial DNA se- 

quences. Evolution 50: 195-221. 

. 1999. The delimitation of phylogenetic spe- 
cies with DNA sequences: A critique of Davis and 
Nixon’s population aggregation analysis. System- 
atic Biology 48: 199-213. 

Brower, A. V. Z. and M. G. Egan. 1997. Cladistics of 
Heliconius butterflies and relatives (Nymphalidae: 
Heliconiiti): The phylogenetic position of Eueides 
based on sequences from mtDNA and a nuclear 
gene. Proceedings of the Royal Society of London 
B 264: 969-977. 

Brown, K. S., Jr. 1979. Ecologia Geografica e Evolu- 


VOLUME 102, NUMBER 3 


¢ao nas Florestas Neotropicais. Universidade Es- 

tadual de Campinas, Campinas, Sao Paulo, Brasil. 

. 1981. The biology of Heliconius and related 
genera. Annual Reviews of Entomology 26: 427— 
456. 

Brown, K. S., Jr., P. M. Sheppard and J. R. G. Turner. 
1974. Quaternary refugia in tropical America: Ev- 
idence from race formation in Heliconius butter- 
flies. Proceedings of the Royal Society of London 
B 187: 369-378. 

Davis, J. I. and K. C. Nixon. 1992. Populations, ge- 
netic variation, and the delimitation of phyloge- 
netic species. Systematic Biology 41: 421—435. 

de Queiroz, K. and M. J. Donoghue. 1988. Phyloge- 
netic systematics and the species problem. Cladis- 
tics 4: 317-338. 

Descimon, H. and J. M. de Maeght. 1984. Semispecies 
relationships between Heliconius erato cyrbia 
Godt. and H. himera Hew. in southwestern Ec- 
uador. Journal of Research on the Lepidoptera 22: 
229-237. 

Doyle, J. J. 1995. The irrelevance of allele tree topol- 
ogies for species delimitation, and a non-topolog- 
ical alternative. Systematic Botany 20: 574—588. 

Eldredge, N. and J. Cracraft. 1980. Phylogenetic Pat- 
terns and the Evolutionary Process. Columbia 
University Press, New York. 

Eltringham, H. 1916. On specific and mimetic rela- 
tionships in the genus Heliconius, L. Transactions 
of the Entomological Society of London 1916: 
101-148. 

Emsley, M. G. 1964. The geographical distribution of 
the color-pattern components of Heliconius erato 
and Heliconius melpomene with genetical evi- 
dence for the systematic relationship between the 
two species. Zoologica NY 49: 245-286. 

Hewitson, W. C. 1854. Illustrations of New Species of 
Exotic Butterflies, V 

. 1867. Descriptions of some new species of 
diurnal Lepidoptera. Transactions of the Entomo- 
logical Society of London, Third Series, 5: 561— 
566. 

Holzinger, H. and R. Holzinger. 1994. Heliconius and 
related genera. Sciences Nat, Venette, France. 
International Commission on Zoological Nomencla- 
ture. 1985. International Code of Zoological No- 
menclature. International Trust for Zoological No- 
menclature and the British Museum (Natural His- 

tory), London. 

Jiggins, C. D., W. O. McMillan, W. Neukirchen, and 
J. Mallet. 1996. What can hybrid zones tell us 
about speciation? The case of Heliconius erato 
and H. himera (Lepidoptera: Nymphalidae). Bio- 
logical Journal of the Linnean Society 59: 221— 
242. 

Kaye, W. J. 1916. A reply to Dr. Eltringham’s paper 
on the genus Heliconius. Transactions of the En- 
tomological Society of London 1916: 149-155. 


685 


Lamas, G. 1998. Comentarios taxonémicos y nomen- 
claturales sobre Heliconiini neotropicales con de- 
signacion de lectotipos y descripcion de cuatro su- 
bespecies nuevas (Lepidoptera: Nymphalidae: 
Heliconiinae). Revista Peruana de Entomologia 
40: 111-125. 

Nixon, K. C. and Q. D. Wheeler. 1990. An amplifi- 
cation of the phylogenetic species concept. Cla- 
distics 6: 211-223. 

Mallet, J. 1993. Speciation, raciation and color pattern 
evolution in Heliconius butterflies: evidence from 
hybrid zones, pp. 226—260. /n Harrison, R. G. ed., 
Hybrid Zones and the Evolutionary Process. Ox- 
ford University Press, Oxford. 

. 1995. A species definition for the Modern 

Synthesis. Trends in Ecology and Evolution 10: 

294-298. 

. 1996. Reply from J. Mallet. Trends in Ecology 
and Evolution 11:174—175. 

Mayr, E. 1940. Speciation phenomena in birds. The 
American Naturalist 74: 249-278. 

. 1963. Animal species and evolution. Belknap 
Press, Cambridge, Massachusetts. 

Mishler, B. D. and M. J. Donoghue. 1982. Species con- 
cepts: A case for pluralism. Systematic Zoology 
31: 491-503. 

Neustetter, H. 1929. Nymphalididae: Subfam. Helicon- 
linae, pp. 1-136. /n Strand, E. ed., Lepidoptero- 
rum Catalogus, part 36. W. Junk, Berlin. 

Riffarth, H. 1902. Nochmals Ch. Oberthiir’s Etudes 
d’Entomologie, Vol. 21. Berliner Entomologische 
Zeitschrift 47: 157-166. 

Sheppard, P. M. 1960. Natural Selection and Heredity. 
Harper Torchbooks, New York. 

Sheppard, P. M., J. R. G. Turner, K. S. Brown, W. W. 
Benson, and M. C. Singer. 1985. Genetics and the 
evolution of Muellerian mimicry in Heliconius 
butterflies. Philosophical Transactions of the Roy- 
al Society of London B 308: 433-613. 

Stichel, H. 1923. Kolombische Heliconius. (Lep., 
Rhop.). Deutsches Entomologische Zeitschrift 
1923: 260-270. 

Turner, J. R. G. 1971. Studies of Miillerian mimicry 
and its evolution in burnet moths and heliconid 
butterflies, pp. 224—260. Jn Creed, R. ed., Ecolog- 
ical Genetics and Evolution. Blackwell Scientific 
Publications, Oxford and Edinburgh. 


APPENDIX | 


Translation of Lamas’ (1998: 119-120) critique. 
Translation is as precise as possible, but some idioms 
were translated freely to improve comprehensibility. 
For complete literature citations, see Lamas (1998) 


Status of Heliconius tristero Brower and H. melpo- 
mene mocoa Brower 

Recently, Brower (1996) has described two new 
taxa of Heliconius from Putumayo, southeastern Co- 
lombia, based on preliminary data from analysis of 


686 


mitochondrial DNA sequences. Both names are dis- 
cussed separately here. 

Heliconius tristero was diagnosed by Brower (1996) 
as a taxon of the species group, belonging to the H. 
cydno Doubleday clade, and narrowly separated from 
H. heurippa Hewitson and H. timareta Hewitson. Al- 
though he did not mention it explicitly, Brower seems 
to imply that heurippa, tristero and timareta could 
constitute a subclade of the cydno group. The three 
taxa exhibit an allopatric distribution, heurippa occur- 
ring in the central eastern region of Colombia, tristero 
in southeastern Colombia and fimareta in eastern Ec- 
uador (and northern Peru, vide infra); while heurippa 
and tristero (so far as is known) are phenotypically 
monomorphic, timareta is polymorphic. Brower (1996: 
330) concluded that “‘... tristero is considered a spe- 
cies only because its geographically adjacent close rel- 
atives, H. heurippa and H. timareta, have been tradi- 
tionally considered species as well.’’ This conclusion 
is not so logical as to be irrefutable. A more valid (and 
refutable) hypothesis would be to consider the three 
taxa conspecific, based on Brower’s own molecular 
and morphological analyses (which would show their 
narrow evolutionary relationship), their allopatric dis- 
tributions (and geographical proximity), and the sup- 
posed existence of transitional forms (“‘hybrids”’) be- 
tween heurippa and tristero (represented by the 
“type” of bellula Stichel, but vide infra), which might 
suggest that, as Mallet (1995) put it nicely in his def- 
inition of species as “‘genotypic aggregations:” “ 
closely related allopatric forms should mostly be con- 
sidered conspecific.”’ Naturally, even though it is well 
known that in the absence of evidence of specific dis- 
tinctness in sympatry the conspecificity of allopatric 
taxa is arbitrary, such arbitrariness is certainly less than 
to assign a taxon to a particular taxonomic level fol- 
lowing a “tradition.” 

In synthesis, there are two opposing taxonomic hy- 
potheses: 1) tristero is a taxon at the species level, 
evolutionarily independent of heurippa and timareta, 
with its own historical destiny; or 2) heurippa, tristero 
and timareta constitute a polytypic species with three 
geographical races (subspecies) which have the same 
historical destiny. With the scarce taxonomic and ge- 
netic information available at the moment, it is impos- 
sible to decide which of these is closer to the truth. 

The case of Heliconius melpomene mocoa seems 
much simpler, and here I offer the hypothesis that mo- 
coa is no more than a new synonym of H. m. bellula 
Turner, 1971, as I will demonstrate below. The name 
bellula was introduced for the first time in the literature 
by Stichel (1923), who proposed it as a form of Hel- 
iconius amaryllis amaryllis C. & R. Felder, thus con- 
stituting an infrasubspecific name (excluded by the 
ICZN). The name bellula recently became available 
when Turner (1971) elevated it to the subspecific level 
(as a subspecies of H. melpomene (L.), making the 
name attributable to Turner (ICZN Arts. 10c, 23j, 50c). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Brower examined the male specimen upon which Sti- 
chel based his infrasubspecific name bellula (and 
which constitutes the holotype of bellula Turner) and 
decided that it represented an individual belonging to 
the cydno clade, and not to melpomene. Brower also 
examined the “‘holotypes” of two other infrasubspe- 
cific names proposed by Stichel (1923), permira and 
degener (which have never been elevated to the sub- 
specific level and thus are nomenclaturally unavail- 
able), suggesting that they might represent hybrids be- 
tween H. heurippa and H. tristero. What is more, he 
assumed that the type of bellula appeared to be a re- 
combinant backcross between these hybrids and tris- 
tero. Arguing that the ICZN “‘forbids”’ the use of spe- 
cies-level names that are based on hybrids, he decided 
to set aside the name bellula and establish new names 
for the member of the cydno clade (tristero) and the 
subspecies of melpomene present in the upper Rio Pu- 
tumayo (mocoa). 

Brower does not seem to have realized that Stichel 
(1923), in addition to bellula, permira and degener, 
described four other “‘forms” of amaryillis from the 
region of the upper Putumayo (anacreontica, perrara, 
rufata and aglaspis), and that other authors introduced 
six additional names for specimens from the same zone 
(parva Neustetter, tenuifasciata Neustetter, aurofascia- 
ta Neustetter, paula Neustetter, paulina Niepelt and 
carminata Niepelt). In my opinion—and contrary to 
Brower’s view—these 13 names pertain to examples 
of melpomene, and represent transitional forms (“‘hy- 
brids’’) between the subspecies bellula Turner and 
malleti Lamas. All those specimens were captured 
around Mocoa (01°09'N, 76°37’W) by collectors who 
worked for Werner Hopp, and were very likely ob- 
tained in the hybrid zone near Villa Garzon (= Villa 
Amazonica; 01°05'N, 76°35’W, 420 m), a site studied 
by Mallet (1993), a few km. East of Mocoa. The ho- 
lotype of bellula is labeled as having been obtained on 
the Rio Guayuyaco (written ““Guagzayaco”’ on the la- 
bel), possibly very near the village of Guayuyaco (= 
Napoles; 01°04'N, 76°26’W). The other specimens 
bear the localities Rio Mulato (01°08'N, 76°36'’W), 
500 m; Mocoa, 530 m; or simple ““Mocoa”’ (for a gen- 
eral description of the area, see Salazar 1995). Mallet 
(1993) referred to the hybrids found near Villa Garz6n 
as the product of the transition between bellula and 
aglaope C. & R. Felder; the correct name of the latter 
subspecies is malleti Lamas, since the subspecies 
aglaope is limited to the lower Rio Maranon, lower 
Rio Huallaga and the Rio Ucayali basin in Peru (La- 
mas 1988). 

Of the 14 names applied to bellula X malleti hy- 
brids, the “‘purest’’ example corresponds to the holo- 
type of bellula, which only shows tiny yellow spots at 
the costal inner edge of the discal red forewing band, 
appearing almost identical to the holotype of mocoa 
(which lacks the yellow spots). Brower based his 
judgement on the presence of these yellow spots, and 


VOLUME 102, NUMBER 3 


the morphology of the genitalia of the bellula holo- 
type, to assert that that specimen was a heurippa X 
tristero “hybrid,” (the same as permira and degener). 
But, in the first place, heurippa (one of the supposed 
parents) is not known from the Mocoa region, only 
from much further north, from the Meta and Guaya- 
bero basins, while surely tristero should occur in the 
basin of the Rio Orteguaza, since bellula has been re- 
ported from there. (It would be very interesting to dis- 
cover if heurippa, or tristero, or both, occur in the Rio 
Caguan basin, between the Guayabero and the Orte- 
guaza). Further, even though Brower admitted that 
“male genitalia of the taxa examined are similar, and 
display substantial intra-racial variability in form,’’ he 
concluded that the genital morphology of the bellula 
holotype corresponded to the cydno clade, based on 
the examination of that one specimen. 

Finally, Brower argued that the name bellula should 
be set aside because it is applied to a hybrid (which, 
as indicated by the discussion above, I consider com- 
pletely erroneous) and because the ICZN “‘forbids”’ the 
use of names applied to hybrids. In the glossary of the 
Code (1985: 256) it is clearly indicated that “‘The 
progeny of two individuals belonging to different sub- 
species of same species are not hybrids.’’ The Code 
does not “forbid” the use of names given to interspe- 
cific hybrids, it simply excludes them from nomencla- 
ture, except for the principle of homonymy. Therefore, 
if the name bellula does not correspond to an inter- 
specific hybrid, its use to designate a subspecific taxon 
is perfectly valid, and in consequence, Heliconius mel- 
pomene mocoa Brower, 1996 is a subjective junior 
synonym of Heliconius melpomene bellula Turner, 
1971. 


APPENDIX 2 


Translation of Stichel’s (1923) description of H. am- 
aryllis amaryllis and H. amaryllis amaryllis f. bellu- 
la.from the upper Putumayo of Colombia. 


687 


H. amaryllis amaryllis Feld. 
Forma typ. 

Two males.—Forewing with a somewhat variable 
broad red discal patch, somewhat like the figure of H. 
amaryllis euryades Riff. in Gen. Ins. v. 112, plate 3, 
fig. 10, but with a more blurred distal margin, and 
which is somewhat indented below the forewing me- 
dian vein. The yellow transverse band of the hindwing 
fragmented near the base by thin black lines, and the 
veins cutting across the band are more or less black. 
Mocoa (Put[umayo]), September, October. 

Among other things, a character of this species is 
said to be the absence of red basal streaks on the costal 
margin of the forewing underside. This feature is pre- 
sent in all of the observed nominate and closely allied 
forms, but based on the other specific characters, par- 
ticularly the position of the hindwing band, they can 
only be considered forms of amaryllis. This band is 
somewhat variable, but it is always positioned such 
that the posterior border lies outside the posterior edge 
of the discal cell. In a third male specimen (Mocoa, 
May), the forewing band is somewhat smaller, so that 
it scarcely touches the end of the cell, the red is faded 
and greasy, which is often seen as a pathological fea- 
ture of red-banded Heliconius. 


Forma bellula f. nov. 

One male.—Most similar to the nominate form, the 
crimson forewing patch more ragged on the margin, 
partly dusted with black scales, the cell almost com- 
pletely free of red, an additional character is a sulfur- 
yellow spot proximal to the subcostal vein. The yellow 
transverse band on the hindwing is broad, extending 
nearly 7 mm. from the apex to the trailing edge. 
Hindwing beneath very similar to H. amaryllis rosina 
Bsd. With red basal streaks on the leading edge of the 
forewing and three red basal spots in the hindwing. 
Forewing length 41 mm. Rio Guaqzayaco 
(Put[umayo]). 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 688-740 


APHID PARASITOIDS (HYMENOPTERA: BRACONIDAE: APHIDITNAE) 
OF NORTHWEST USA 


K. S. PIKE, P. STARY, T. MILLER, G. GRAF, D. ALLISON, L. BOYDSTON, AND R. MILLER 


(KSP) Entomologist and (DA, LB, GG) Research Technicians, Washington State Uni- 
versity, Irrigated Agriculture Research & Extension Center, 24106 N Bunn Road, Prosser, 
WA 99350, U.S.A.; (PS) Entomologist, Institute of Entomology, Academy of Sciences of 
the Czech Republic, BraniSovska 31, 370 05 Ceské Budéjovicé, Czech Republic; (RM) 
Entomologist, University of Guam, CALS, AES, 303 University Drive, UOG Station, 
Mangilao, GU 96923, U.S.A.; (TM) Manager, Northwest Biological Control Insectary & 
Quarantine, Washington State University, Pullman, WA 99164, U.S.A. 


Abstract.—A comprehensive assessment is provided of the Northwest USA aphid-par- 
asitoid species (Hymenoptera, Braconidae, Aphidiinae). Eighty-one species of parasitoids 
in 19 genera are recognized; 35 species are new to the region; 7 species are newly 
described: Aphidius segmentatus Pike and Stary [hosts: Hyperomyzus on Hieracium, Lac- 
tuca, Senecio, and other Compositae, and on Uroleucon on Achillea]; Betuloxys alnicolus 
Pike & Stary [host: uncertain, possibly Boernerina variablilis Richards on Alnus]; Mon- 
octonus campbellianus Pike & Stary [hosts: Brachycaudus, Hyalopterus, and Phorodon 
on Prunus]; Monoctonus pacificus Pike & Stary [host: Macrosiphum tuberculaceps (Es- 
sig) on Achlys triphylla|; Praon coniforme Pike & Stary [host: Aphthargelia symphori- 
carpi (Thomas) on Symphoricarpos];,; Praon fulvum Pike & Stary [hosts: Macrosiphum 
spp. on Achlys, Crepis, Lupinus, Spiraea]; Trioxys setaceus Pike & Stary [host: Acyrtho- 
siphon macrosiphum (Wilson) on Amelanchier]. Seventy genera of aphids (200+ spp.) 
were attacked by aphidiine parasitoids. Some 400+ parasitoid-aphid associations, and 
hundreds more tritrophic (parasitoid-aphid-plant) combinations are reported for the first 
time in North America. 


Key Words: aphid, parasitoid, systematics, Northwest USA 


The aphidiine parasitoids of Northwest 
USA were previously understood to include 
about 54 species in 13 genera; these were 
linked with about 100 species of aphids 
(Pike et al. 1996, 1997, 1999). Based on 
new environmental studies, the parasitoid 
guilds are recognized now to be much larg- 
er. Presented here is a comprehensive re- 
view of Northwest species, including an 
abridgment of previously published records. 

Eighty-one species of aphidiine parasit- 
oids in 19 genera are now recognized in 


Northwest USA; 35 species are new to the 
region; 7 species are newly described. Col- 
lectively, these parasitoids attacked more 
than 200 species in 71 genera of aphids. 
Some 400+ parasitoid-aphid associations, 
and hundreds more tritrophic (parasitoid- 
aphid-plant) combinations are reported for 
the first time in North America. 

The work is a foundation study of the 
Northwest aphidiine fauna. It provides 
background documentation for possible fu- 
ture studies on a variety of fronts, such as 


VOLUME 102, NUMBER 3 


Table 1. Legend for listings. 


689 


PARASITOID 


Praon americanum (Ashmead) 


[Example] 


Aphis coweni Palmer: WA, KITTITAS [*] — Stampede Pass, 8-VIII-96 on Veratrum viride (96K1477f). 


APHID HOST STATE COUNTY SPECIFIC DATE APHID PLANT WSU CODE 
COLLECTION COLLECTED Host 
SITE 


[*] indicates previous published record, see Pike et al. 1996, 1997, 1999. 

[+] indicates collection contained more than one species of aphid. 

[+] indicates parasitoid-aphid association shown is not certain. 

Abbreviations: ALE, Arid Lands Ecology Reserve; CA, California; CG, Campground; Cr, Creek; det., deter- 
mined; E, east; FR, forest road; G&P, Gillette & Palmer; HRL, Hille Ris Lambers; HMU, Habitat Management 
Unit; ID, Idaho; Lk, Lake; mi, mile; MT, Montana; Mt, Mount; Mtn, Mountain; N, north; nr, near; OR, Oregon; 
RNP, Mount Rainier National Park; Pk, Park; Rd, Road; S, south; sp., species; SP, State Park; W, west; WA, 
Washington; WLA, Wildlife Area or Refuge; WSU, Washington State University; YIR, Yakama Indian Reser- 


vation. 


biosystematics, including new species de- 
scriptions; parasitoid guilds; population ge- 
netics; host adaptation, switching, and cross 
habitat movement; geographic spread of ex- 
otic species; interspecies population devel- 
opment; ecosystem relationships and effects 
on parasitoid and predator complexes, and 
target pests; application and advantages of 
aphid-plant biodiversity for maintaining, 
stabilizing, and/or increasing parasitoids of 
merit; and uses and possible export of 
Northwest parasitoids to other global areas. 


MATERIALS AND METHODS 


Aphidiine parasitoids were obtained from 
aphid rearings, with the aphids collected 
from a wide range of host plants from 
Northwestern USA, principally Washing- 
ton. In total, more than 5,300 aphid collec- 
tions were taken, of which 2,940 were par- 
asitized. The aphid populations sampled 
varied in size. Where possible 50 to >200 
aphids were taken per sample; a subsample 
of 5 to 30 aphids were preserved in 70% 
ethanol for later identification. For the par- 
asitoid rearings, aphids were held in-labo- 


ratory at 20 + 3°C for 25-30 days in 
screen-covered semi-transparent plastic 
containers (two sizes used: 300 ml, 10 cm 
dia X 4 cm ht; and 3500 ml, 19 cm dia X 
13 cm ht) or occasionally in paper lunch 
bags. After emergence, the parasitoids were 
placed in 70% ethanol for subsequent de- 
termination. Voucher specimens of both 
aphids and parasitoids are in Washington 
State University-Prosser collections, Pros- 
ser, WA. 

Descriptions of new species were based 
on whole dry and dissected slide-mounted 
specimens examined under 40—400X mag- 
nification. Body lengths were measured and 
recorded in millimeters. Holotypes were 
dry-mounted on paper tabs and pinned. De- 
scriptive terminology is after Huber and 
Sharkey (1993). Holotypes are deposited in 
the National Museum of Natural History, 
Smithsonian Institution, Washington, DC 
(USNM). 

For cross reference and other information 
on aphid parasitoids of North America, see 
Gahan 1911; Smith 1944; Liu 1977; Marsh 
1979; Johnson 1987; Pike et al. 1996, 1997, 


690 


1999; for world information, see Mackauer 
and Stary 1967. 

Aphid nomenclature follows Remaudiére 
and Remaudiére 1997. 


RESULTS 


Aphidiine parasitoid genera and _ spe- 
cies.—Parasitoid genera and species are 
listed alphabetically, as are aphid hosts un- 
der each species. Locations are indicated by 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


state (large caps), county (small caps), and 
nearest city or identifiable site. Numbers in 
parentheses represent authors’ codes of 
specimens (aphid-parasitoid vouchers) in 
Washington State University—Prosser col- 
lections (Table 1). Collections and deter- 
minations were by authors unless otherwise 
specified. Aphids collected by A. Jensen 
were determined by A. Jensen. Authorship 
of new species is attributed to K. S. Pike 
and P. Stary. 


APHIDIINE PARASITOIDS OF NORTHWEST USA 


Genus ACANTHOCAUDUS Smith 


Acanthocaudus caudacanthus (Smith) 
Uroleucon sp.: WA, PEND OREILLE—Le Clerc WLA, 3-VIII-95 on Aster sp. (95T400); SPOKANE—nr Peone, 
24-VIII-95 on Centaurea pratensis (95T473*). 


Acanthocaudus sp.? 
Undetermined sp.: WA, GARFIELD—Pataha Cr, nr FR160 & FR40, 21-[X-95 on Chimaphila umbellata 
(95T597). 


Genus ADIALYTUS Foerster 


Adialytus ambiguus (Haliday) 
Sipha elegans del Guercio: MT, SANDERS [on Agropyron intermedium*]. 
Sipha sp.: WA, ASOTIN [on unknown Gramineae* ]. 


Adialytus fuscicornis (Ashmead) 

Aphis armoraciae Cowen: OR, UMATILLA—Hermiston, 17-VI-90 on unknown plant (90J002); WA, YAKIMA— 
Wenas Lk, 30-V-96 on Erysimum arenicola (96G120), & 11-VI-97 on Lomatium sp. (97G051*). 

Aphis helianthi Monell: WA, YAKIMA—Wenas Lk, 11-VI-97 on Lomatium sp. (97GO0517¢). 

Forda marginata Koch: WA, GARFIELD—Dye Seed Airfield, 5-[X-95 on Bromus sp. (95G542); Kirby, 1-[X- 
95 on Bromus inermis (95K160). 

Undetermined sp.: WA, AsoTIN [on Populus trichocarpa*}. 

Adialytus salicaphis (Fitch) 

Chaitophorus populicola Thomas [on Populus trichocarpa unless otherwise indicated]: ID, BOUNDARY [*]; 
WA, AsotTin—Asotin WLA, 31-VII-96 & 15-VIII-96 (96T355, 96T454); BENTON—Grandview, 10-X-95 
(95G694); WHITMAN—Pullman, 9-X-96 on Populus tremuloides (96T582); YAKIMA—YIR, Lucy Canyon, 
25-IV-96 & 16-V-96 (96G021, 96G061) & nr Satus, 1-VI-96 (96K062). 

Chaitophorus populifolii (Essig) [on Populus trichocarpa]: WA, KitrirAs—Manastash Cr, 16-VIII-96 
(96G485); Ellensburg, 13-VI-97 (97G083); YAKIMA—Naches, 27-IX-96 (96G600). 

Chaitophorus utahensis (Knowlton) [on Salix sp.]: WA, AsoTiN—Chief Timothy HMU, 16-VIII-95 (95T442); 
BENTON—nr Grandview, 24-VIII-95 (95G514); YAKIMA [*]—Grandview, 22-V-95 & 15-V-96 (95G078, 
96G047). 

Chaitophorus viminalis Monell [or nr viminalis] [on Salix sp.]: WA, OKANOGAN—Alta Lk, 10-L[X-97 (97K026, 
97K028); YAKIMA—Wenas Cr, 13-VIII-96, (96G459). 

Chaitophorus sp. {nr nigrae Oestlund & pallipes Richards]: WA, AsotiN—Heller’s Bar, 11-VII-95 on Salix 
sp. (95T2107). 

Chaitophorus sp. {on Salix sp. unless otherwise indicated]: WA, AsoTIN [*]; BENTON [*]—Prosser, 10-X-95 
on Populus trichocarpa (95G693); GRANT—Potholes Reservoir, 13-VI-97 (97G101). 

Periphyllus nr brevispinosus G&P: WA, PEND OREILLE—Sullivan Lk Rd, Harvey Cr, 3-VIII-95 on Acer sp. 
(951395); SPOKANE—Mt Spokane SP, 24-VIII-95 on Acer sp. (95T487). 

Periphyllus sp.: WA, ASOTIN [on Acer sp. *]. 


VOLUME 102, NUMBER 3 691 


Genus APHIDIUS Nees 


Aphidius avenaphis (Fitch) 

Acrthosiphon lactucae (Passerini): WA, WHITMAN—Pullman, 13-VI-96 on Lactuca serriola (96T0327#). 

Diuraphis noxia (Kurdjumov): WA, AsoTin—Anatone, 24-VII-96 & 22-VII-97 on Hordeum vulgare (961596, 
97JOO5—97J023); KLICKITAT—Bickleton, 29-VI-97 & 6-VIII-97 on Triticum aestivum (97G300, 97G354). 

Rhopalosiphum padi (L.): WA, GRANT—Ephrata, 8-VII-97 on Zea mays, K. Volker collector (97K058). 

Sitobion avenae (FE) [on Triticum aestivum unless otherwise indicated]: ID, LArAH—Moscow Mtn, 3-VIII-96 
on unknown Gramineae (9613967); SHOSHONE—Thompson Pass, 29-VII-95 on Agropyron caninum 
(95G478), Arabis sp. (95G475+), & Poa nervosa (95G470); OR, UMATILLA—Harris Pk, 11-VI-96, on 
Dactylis glomerata (96G230+); WA, ADAMsS—Bruce, I-VII-96 (96B004, 96B007); Othello, 26-VI-95, 1- 
VII-96 (95B009, 96B009); AsoTin—Anatone, 24-VII-96 on Hordeum vulgare (961337); BENTON—Prosser, 
2, 28 & 30-VI-95, 17-VII-95, 24-VI-96, 1 & 8-VII-96, 3-VII-97, (95G103, 95G276, 95G279, 95G285, 
95G352, 95G354, 95G357+, 95G358, 96G271, 96G312, 96G328, 96G330, 97F201): Richland, 21-VI-95 
(95G2217+); West Richland, 28-VI-95 (95G279); WSU-Prosser, 18-VII-97 on Setaria lutescens (97G243); 
DouGLAs—Waterville, 24-VII-96 (96B025); FRANKLIN—Kahlotus, 6-VII-95 (95G290, 95G3087); Star 
School, 6-VII-95 (96G289); GRANT—WSU-Royal Slope, 26-VI-95 (95B007); Kittiras—S. Fork 
Manastash Cr, 24-VII-95 on Agropyron repens (95G375), Agropyron caninum (95G383), & T. aestivum 
(95G372); 11-IX-95 on Bromus sp. (95G384); Manastash Rd, 11-IX-95, on D. glomerata (95G565); 
Stampede Pass, 8-VIII-96 on Phleum sp. (96K142); KLickiTaT [on H. vulgare & Triticosecale rimpani* | 
—Bickleton, 9 & 17-VII-95 (95G317+—95G3197, 95G362), 7-VIH-97 (97G224); Trout Lk, 29-VI-97 on D. 
glomerata (97G160); LINcoLN—Harrington, 19-VII-96 (96B023); Davenport, 16 & i9 -VII-96 (96B011, 
96B020, 96B023); Reardan, 16-VII-96 (96B014); PlERCcE—Graham, 25-V-96 on unknown grass (96T001); 
WHITMAN—Colton, 31-VII-96 on H. vulgare (96T358A); Hay, 6-VII-95 (95G307); Pullman, 5-VII-94, 19- 
VII-96 (94T022, 96T320A); Ridpath, 6-VII-95 (95G299+, 95G3007); Rosalia, 16-VII-96 (961294); 
YAKIMA—Sunnyside, 28-VI-95 & 5-VII-95 (95G272, 95G286); Zillah, 28-VI-95 (95G270); 25-VI-95 on 
D. glomerata (95G250); YIR, nr Signal Peak, 25-VI-95, on Agropyron spicatum (95G416); Chinook Pass, 
24-VII-97, on Poa gracillima (97G260); 24-VII-97, on Poa nervosa (97G268). 

Undetermined spp.: WA, GARFIELD—Ruckert Rd & Fitzgerald Rd, 15-VII-96 on Hordeum sp., W. Turner & 
J. Rumph collectors (96J069A); KinGc—Stampede Pass, 8-VIII-96 on Caltha biflora (96G447); PEND 
OREILLE—Le Clerc WLA, 3-VIII-95 on unknown grass (95T401). 


Aphidius colemani Viereck 
Aphis nerii Boyer de Fonscolombe: WA, YAKIMA—Union Gap, 23-X-95 on Asclepias speciosa (95G725). 
Brachycaudus helichrysi (Kaltenbach): WA, WHITMAN—Pullman, 28-VI-96 on Anthemis arvensis (96T 162). 
Hyadaphis foeniculi (Passerini): WA, ISLAND—Clinton, 4-VI-96 on Lonicera sp. (96G156). 
Lipaphis erysimi (Kaltenbach): WA, WHITMAN—Pullman, 3-VI-96 on unknown plant (96T 1637). 
Macrosiphum euphorbiae (Thomas): WA, WHITMAN—Pullman, 3-VII-96 on unknown plant (96T1637). 
Myzus persicae (Sulzer): WA, SNOHOMISH—Monroe, 24-VII-96 on Pisum sativum (9615947). 
Myzus sp.: WA, SPOKANE—Spokane, 11-VII-96 on Antirrhinum majus (96T258); WHITMAN—Pullman, 14-IX- 

93, on Forsythia (93T019). 

Undetermined sp.: WA, KING [on Hibiscus sp.*]. 


Aphidius ervi Haliday 

Acyrthosiphon kondoi Shinji: 1D, SHOSHONE—Thompson Pass, 29-VII-95 on Melilotus alba (95G4667); WA, 
KLICKITAT, Hwy 14 nr Alderdale, 14-V-96 on Melilotus sp. (96G0337). 

Acyrthosiphon lactucae (Passerini) [on Lactuca_ serriola]: ID, BENEWAH—McCroskey SP, 31-VIII-95 
(95T514+); WA, AsoTiIn—Anatone, 28-VI-96 (96T088); Asotin, 28-VI-96 (96T068); Asotin Cr WLA, 5- 
VII-96 & 8-VIII-96 (96T 144, 96T419); Couse Cr, 10-VIII-95 (95T422); Snake River Rd, 18-VII-95 & 28- 
VI-96 (95T249+, 96T072); South Fork Asotin Cr, 22-VII-95 & 11-VII-96 (95T468, 96T230); BENTON— 
Prosser, 17-VII-95 (95G359+); 24-VI-96 & 8-VIII-96 (96G270, 96G434); ALE, 19-V-95 (95G071); 
CHELAN—Chelan SP, 11-[X-97 (97K038); SPOKANE—Plaza, 16-IV-96 (96T279); WHITMAN—Pullman, 16, 
30-VII-96 & 1, 8-VIII-96 (961304, 961362, 96T382, 961436); Ridpath, 6-VII-95 (95G296); YAKIMA— 
Mabton, 28-VI-95 (95G265); Sunnyside, 22-V-95 (95G076); Wenas Lk, 11-VI-97 (97G055). 

Acyrthosiphon pisum (Harris) [on Medicago sativa unless otherwise indicated]: ID, SHOSHONE—Thompson 
Pass, 29-VII-95 on Melilotus alba (95G466+); MT, SANDERS [on Lathyrus odoratus*|; WA, ADAMS— 
Macmaranon Rd, 2-VIII-96 (96B026); WSU-Othello, 26-VI-95 (95B008); AsoTtiIn—Couse Cr & Snake 
River Rds, 18-VII-95 on Helianthus annuus (95T252); BENTON [on M. sativa & Trifolium pratense*| — 
28-VI-95 (5G277+); Prosser, 27-VI-96 on Vicia angustifolia (96A009); 23-VII-9, on T. pratense (96G391); 
WSU-Prosser, 26, 28-IV-97 & 14-V-97 (97G014, 97G015, 97G030); Whitstran, 1-V-97 (97G019); 


692 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Co_LuMBIA—Hwy 124 & Tucannon Rd, 5-X-95 (95G678+); GRANT—WSU-Royal Slope, 26-VI-95 
(95B003); IsLAND—South Whidbey, 4-VI-96 on Trifolium sp. (96G161); KLICKITAT—Hwy 14 nr Alderdale, 
14-V-96 on Melilotus sp. (96G0337+); Roosevelt, 24-IV-96, on Melilotus sp. (96GO147+); LINCOLN—Lk 
Roosevelt, 31-VII-97 on M. lupulina (97G304); SNoHOMISsH—Monroe, 24-VII-96 on Pisum sativum 
(96T594+); SPOKANE—Four Lk, 25-VII-97 on Allium sp. (97KO005); WHITMAN—Pullman, 16, 30-VII-96 on 
P. sativum (961303, 96T359); 14-IV-96 on Trifolium sp. (96T272); 26-VI-96 on Vicia sp. (96T063) & 
unknown legume (96T064), & 5-VII-96 on Lathyrus oderata (96T178); Ridpath, 6-VII-95 on Lactuca 
serriola (95G296), Medicago sativa (95G297), & Vicia sp. (95G302); YAKIMA—Outlook, 1-VI-96 
(96K059); Rimrock Lk, 23-VU-96 on Melilotus sp. (96G383); Wenas Valley, 30-V-96 (96G111); YIR, 
Signal Peak, 25-VII-95 on Vicia sp. (95G408) & 19-IX-95 on Melilotus sp. (95G625*). 

Aphis craccivora Koch: WA, WHITMAN—Pullman, 25-VII-95 on Capsella bursa-pastoris (951331); 14-1V- 
96, on Trifolium sp. (96T271). 

Aphis sp.: ID, LATAH [on Solanum Lycopersicon *|; WA, ASOTIN—De Spain Springs, 1 1-VI-96 on Umbelliferae 
sp. (96T254). 

Aspidophorodon longicauda (Richards): WA, SKAMANIA—South Prairie, 11-VU-95 on Spiraea sp. (95G343). 

Brachycaudus helichrysi (Kaltenbach) [on Amsinckia sp.]: WA, AsoTiIN—Anatone, 28-VI-96 (96T089); 
FRANKLIN—Eureka, 14-V-96 (96G041); SkAGIT—Campbell Lk, 4-VI-96 on Prunus sp. (96G1657). 

Brevicoryne brassicae (L.): WA, BENTON [on Brassica napus*). 

Chaetosiphon nr hottesi Stroyan: WA, CLARK—Vancouver, 14-I[X-95 on Fragaria sp. (95VO002). 

Diuraphis noxia (Kurdjumoy) [on Triticum aestivum unless otherwise indicated]: WA, AsoTIN [on Avena 
fatua*| AsoTin—Anatone, 22-VII-97 on Hordeum vulgare (97JO007—97J026); BENTON [on H. vulgare & T. 
aestivum* |—Wsvu-Prosser, 28-V-96 (on H. vulgare, 96G087, 96G088), 24-VII-97 (97G274); KLICKITAT— 
Rock Cr, 30-V1-95 (95G283); Bickleton, 7-VII-97 (97G223, 97G238); Cleveland, 7-VII-97 (97G232, 
97G234); WALLA WALLA—Hadley, 11-VI-96 on H. vulgare (96G240); WHITMAN—Pullman, 9-VII-96 on 
H. vulgare, W. Turner & J. Rumph collectors (96J053—96J055); YAKIMA [*]. 

Hayhurstia atriplicis (L.) [on Chenopodium alba\: WA, BENTON—-WSU-Prosser, 18 & 25-VII-97 (97F238, 
97F262); WHITMAN—Pullman, 16-VII-96 (96T302). 

Hyalopteroides humilis (Walker) [on Dactylis glomerata|: WA, SKAGIT—Bow, 5-V-96 (96K026); Edison, 5- 
V-96 (96K027). 

Hyalopterus pruni (Geoffroy): WA, SkKAGIT—Campbell Lk, 4-VI-96 on Prunus sp. (96G1657#). 

Hyperomyzus lactucae (L.): WA, YAKIMA—Yakima Arboretum, 26-IX-95 on Sonchus oleraceus (95G6587). 

Hyperomyzus nigricornis (Knowlton): ID, BENEWAH—McCroskey SP, 31-VII-95 on Lactuca_ serriola 
(95T514+); WA, GARFIELD—Stentz Springs, 22-VII-95 on Senecio sp. (95T456); YAKIMA—YIR, Cedar Cr, 
19-IX-95 on Agoseris sp. (95G622). 

Illinoia davidsoni (Mason): WA, KLickirAat—Trout Lk, 11-VH-95 on Rubus parviflorus (95K069). 

Illinoia spp.: WA, Kirtrras—Quartz Mtn, 25-VII-96 on Rhododendron albiflorum (96G405); PIERCE—RNP, 
Tipsoo Lk, 16-VIII-95 on Spiraea densiflora (95A037); SkAcit—La Conner, 5-VI-96 on Rhododendron 
sp. (96G193); SNoHOMISH—Monroe, 5-VI-96 on Chaenomeles sp. (96G201). 

Macrosiphum albifrons Essig: WA, PIERCE—RNP, Tipsoo Lk, 11-[X-96 on Lupinus sp. (96G547). 

Macrosiphum euphorbiae (Thomas): ID, LATAH—Moscow, 7-X-96 on Solanum sp. (961577); WA, ASOTIN— 
De Spain Springs 11-VI-96 on Potentilla sp. (96T250), 1-VII-95 on Senecio sp. (95T381), 5 & 11-VII- 
96 on Compositae (96T159, 961T249+); BENTON—Prosser, 15-V-96 on Tulipa sp. (96G045); 24-X-96 on 
Medicago sativa (96G651); WSU-Prosser, 27-X-95 on Lamium amplexicaule (95K2137), 18-XI-95 on 
Brassica napus (95G751+), & 1-X-97 on Malva neglecta (97G471+); GARFIELD—nr FR40 & FR44, 22- 
VIII-95 on Osmorhiza chilensis (95T464+); SKAMANIA—Twin Falls CG, 4-VIII-95 on Potentilla sp. 
(95K098+); YAKIMA—Bird Cr Meadows, 26-VI-95, on Castilleja (95A017+); YIR, Old Maid Canyon, 25- 
VI-95 on Cynoglossum grande (95G440). 

Macrosiphum gaurae (Williams): WA, AsoTin—Anatone, 28-VI-96 on Oenothera sp. (96T073). 

Macrosiphum nr pallidum (Oestlund): WA, SKAMANIA—Twin Falls CG, 4-VIII-95 on Potentilla sp. 
(95KO098+74). 

Macrosiphum rosae (L.): WA, SNOHOMISH—Snohomish, 5-V1—96 on cultivated Rosa sp. (96G198). 

Metopolophium dirhodum (Walker): OR, UMATILLA—Harris Pk, 11-VI-96 on Elymus glaucus (96G221); WA, 
KLIckITAT—John Day Dam, 14-V-96 on Phalaris arundinacea (96G036+); LiINCOLN—Lk Roosevelt, 
Goldsmith CG, 31-VII-97 on P. arundinacea (97G305). 

Myzus nr ornatus Laing: WA, SkAGir—La Conner, 5-VI-96 on Trifolium pratense (96G1957+£). 

Myzus persicae (Sulzer): WA, SkAGir—La Conner, 5-VI-96 on Trifolium pratense (96G1957+); BENTON— 
WSU-Prosser, 1-X-97 on Solanum tuberosum (97G477+); WHItMAN—Pullman, 16-VII-95 on Raphanus 
sativum (951248); 8-VIII-95, on Solanum nigrum (95T412); 31-VII-96 on Beta vulgaris (96T366). 


VOLUME 102, NUMBER 3 693 


Myzus sp.: WA, WHITMAN—Pullman, 14-IX-93 on Forsythia sp. (93TO19). 

Ovatus crataegarius (Walker): WA, GRANT—Othello, 12-VII-95 on Mentha piperita (95BO10). 

Phorodon humuli (Schrank): WA, SkAGIt—Campbell Lk, 4-VI-96 on Prunus sp. (96G1657+). 

Rhopalosiphum maidis (Fitch): WA, BENTON—-WSU-Prosser, 18-XI-95 on Capsella bursa-pastoris (95G7507). 

Rhopalosiphum padi (L.): WA, AsoTiN—De Spain Springs, 25-VII-96 on Aquilegia sp. (9613477); BENTON 
[on Triticum aestivum*]|; DouGLAS—Waterville, 12-VIII-95 on 7. aestivum (95B016); FRANKLIN—nr 
Kahlotus, 26-X-95 on T. aestivum (95K2077+); GRANT—Ephrata, 8-VII-97 on Zea mays (97K058); WALLA 
WALLA—Prescott, 30-X-97 on Phalaris arundinacea (97G516). 

Sitobion avenae (FE) [on Triticum aestivum unless otherwise indicated]: ID, BOUNDARY—Moyie Springs, 29- 
VII-95, on Avena sativa (95G445); SHOSHONE—Thompson Pass, 29-VII-95 on Poa nervosa (95G470) & 
Arabis sp. (95G475+); WA, ADAMs—Othello, 1-VII-96 (96B004, 96B007), WSU-Othello, 26-VI-95, 1-VI- 
96 (95B009, 96B009); Asotin [*]—Clarkston, 18-VII-95 on Gramineae (95T255); Field Springs SP, 5- 
VIU-93 on Gramineae (93T013); Couse Cr, 18-VII-95 on Setaria sp. (95T259); Anatone, 12-VII-94 
(94T023); BENTON [on Hordeum vulgare & T. aestivum* |—Prosser, 25, 28 & 30-VI-95, (95G253, 95G274, 
95G276, 95G285); 17-VII-95 (95G352, 95G354, 95G355, 95G358); 24-VI-96 (96G271); 1 & 8-VII-96 
(96G312 & 96G328); WSU-Prosser, 20-I-96 on Capsella bursa-pastoris (96G003+); 18-VII-97 on Setaria 
lutescens (97G243); Richland, 28-VI-95 (95G279); CoLuMBIA—Dayton, 16-VI-94 (94T020); GRANT— 
WSU-Royal Slope, 26-VI-95 (95B007); FRANKLIN—Pasco, Star School, 6-VII-95 (95G289); KLICKITAT— 
Bickleton, 9-VII-95 (95G318): Cemetery Rd nr Bickleton, 7-VII-97 on Hordeum vulgare (97G2267); 
KitrirAs—S. Fork Manastash Cr, 24-VII-9, on Agropyron caninum (95G383); Ellensburg, 9-VII-96 
(96G331); Buck Meadows, 9-VII-96 on Dactylis glomerata (96G346); Stampede Pass, 8-VII-96 on Phleum 
sp. (96K142); LINcoLN—Almira, 26-VI-95 (95B002); Davenport, 16 & 19-VII-96 (96B011, 96B020); 
Harrington, 19-VII-96 (96B023); Reardan, 16-VIH-96 (96B014); Wilbur, 19-VII-96 (96B019); PrERCE— 
Graham, 25-V-96 on Gramineae (96TO01); RNP, Tipsoo Lk, 11-IX-96 on Poa sp. (96G543); WHITMAN 
[*]—Hay, 6-VII-95 (95G307); Pullman, 5-VII-94 (94T022); Colton, 31-VII-96 on H. vulgare (96T358A); 
Rosalia, 16-VII-96 (96T294); YAkIMA—Grandview, 5-VuI-95 (95G286); 23-V-96 on D. glomerata 
(96G074); Zillah, 28-VI-95 (95G270, 95G272); 28-VI-95 on Avena sativa (95G271). 

Uroleucon (Lambersius) sp.: WA, KirrirAs—Lost Lk Trailhead, 6-VIUI-97 on Achillea millefolium (97G318). 

Undetermined spp.: WA, BENTON—Prosser, 25-X-95 on Berberis aquifolium (95G732); 29-X-95 on Senecio 
vulgaris (95G735); CHELAN—Stevens Pass, 16-IX-95 on Gramineae (95K137); KLICKITAT—Cleveland, 7- 
VII-97 on Lotus pinnatus (97G233); WHITMAN—Pullman, 1-VII-96 on Penstemon sp. (96T125), 5 & 9- 
VII-96 on Geranium viscosissinum (961171, 961228); 9-VII-96 on Vicia sp. (96T229). 


Aphidius nr hortensis Marshall 
Ericaphis gentneri (Mason): WA, CHELAN—Stevens Pass, 16-[X-95 on Spiraea densiflora (95K138). 
Liosomaphis berberidis (Kaltenbach): OR MULTNOMAH & WA, YAKIMA [on Berberis sp.*]. 


Aphidius kakimiaphidis Smith 

Nasonovia alpina (G&P): WA, YAKIMA [on Mimulus lewisii*]. 

Nasonovia aquilegiae (Essig) [on Aquilegia formosa or sp.]: WA, BENTON—Prosser, 24-VI-96 (96G278); 
Kittitas [+*]—Buck Meadows, 16-VIII-96 (96G501); YAKIMA—Yakima Arboretum, 3-VI-96 (96G249); 
YIR, Old Mead Canyon, 25-VII-95 (95GG441). 

Nasonovia cynosbati (Oestlund): WA, Kitriras—Lost Lk, 7-VI-94 on Ribes sp. (94K026). 

Nasonovia nr cynosbati (Oestlund): WA, YAKIMA—YIR, nr Mt. Adams Lk, 4-VIII-95 on Ribes viscosissimum 
(95K089). 

Nasonovia nr houghtonensis (Palmer): WA, YAKIMA—Nile, 28-VII-97 on Collomia linearis (97G282). 

Nasonovia wahinkae (Hottes): WA, Kittiras—Buck Meadows, 11-IX-95 on Aconitum columbianum 
(95GS557). 

Nasonovia sp.: MT, SANDERS [on Castilleja sp.*]; WA, Lewis—RNP, Stevens Canyon, 23-VII-96 on 
Pedicularis sp. (96G3727). 


Aphidius lupini Liu 

Macrosiphum albifrons Essig [on Lupinus sp.]: WA, BENTON—Prosser 22-VI-95 & 24-VI-96 (95G249, 
96G276); Kittrras—S. Fork Manastash Cr, 24-VII-95 (95G390); Quartz Mtn 25-VII-96 (96G408); 
PIERCE—RNP, Tipsoo Lk, 11-[X-96, 5-IX-97 (96G547, 97G362); WHITMAN—Pullman, 1-VII-96 (96T129); 
YAKIMA [*]—YIR, nr Howard Lk, 19-[X-95 (95G638) & nr Mt Adams Lk, 25-VII-95 (95G419): Little 
Naches River, nr Timothy Meadows, 9-IX-97 (97G389). 

Aphidius matricariae Haliday 

Aphis fabae Scopoli [on Cirsium arvense]: WA, AsoTin—Sangster Rd, 22-VII-97 (97J003++, Turner & Rumph 
collectors); KitrrrAs—Buck Meadows, 11-IX-95 (95G580). 


694 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Aphis helianthi Monell: WA, YAKIMA—Chinook Pass Hwy & Bald Mt Rd, 23-X-97 on Cirsium arvense 
(97G499). 

Aphthargelia symphoricarpi (Thomas): WA, Kittiras—Lost Lk, 6-VIII-97 on Silene douglasii (97G345). 

Brachycaudus cardui (L.): WA, ASoTIN—Sangster Rd, 22-VII-97 on Cirsium arvense, Turner & Rumph 
collectors (97J0037). 

Brachycaudus helichrysi (Kaltenbach): WA, BENTON—WSU-Prosser, 1-X-97 on Solanum tuberosum 
(97G478); KinG [on Senecio vulgarais*]; KLIcKirAt—Bickleton, 30-V1—95 on Chrysanthemum sp. 
(95G282); WHITMAN—Pullman, 26-VI-96, on Achillea millefolium (96T065). 

Brevicoryne brassicae (L): WA, BENTON—-WSU-Prosser, 6-VII-95 on Brassica napus (95A024). 

Capitophorus elaeagni (del Guercio) [on Cirsium arvense or sp.]: WA, Kittiras—Buck Meadows, 11-IX-97 
(97G437+); YAKIMA—Mabton, Morgan Lk, 28-VI-95 (95G266); Little Naches River (FR 789 & FR 1914), 
9-IX-97 (97G409); 9-IX-97 (97G408); Little Naches River, nr Timothy Meadows, 9-IX-97 (97G394). 

Diuraphis noxia (Kurdjumov) [on Triticum aestivum]: WA, BENTON—Rattlesnake Hills, 17-VII-95 (95G3577); 
WSU-Prosser, 30-IX-97 (97F457+, 97463+); YAKIMA—Mabton Grade, 31-X-97 (97G5177). 

Dysaphis plantaginea (Passerini): WA, CHELAN—-WSU-Wenatchee, 30-V-96 on Pyrus malus (96RO117#). 

Hyalopterus pruni (Geoffroy): WA, SkKAGIT—Campbell Lk, 4-VI-96 on Prunus sp. (96G1657+). 

Illinoia corylina (Davidson) [det. by A. Jensen]: WA, WHITMAN—Pullman, 14-VIII-96 on Aquilegia sp. 
(96T442). 

Macrosiphum euphorbiae (Thomas): WA, BENTON—-WSU-Prosser, 1-X-97 on Malva neglecta (97G4717). 

Myzus persicae (Sulzer): ID, BONNER—Sandpoint, 30-IV-97 on Viola tricolor (97T006); OR, UMATILLA— 
Hermiston, 23-VII-93 on unknown plant, A. Jensen collector (93J005); WA, BENTON—WSU-Prosser, 27- 
X-95 on Malva neglecta (95K212+), 1-X-97 on Chenopodium album (97G470*), Solanum sarrachoides 
(97G4721), & S. tuberosum (97G476); KLickiIrat—John Day Dam, 28-IV-95 on Veronica anagallis- 
aquatica (95G009%+); SPOKANE—Spokane, Manito Pk, 19-IX-96 on Syngonium sp. (96T538); WHITMAN— 
Pullman, 22-VII-95 on Solanum lycopersicon (9512997). 

Myzus sp.: WA, WHITMAN—Pullman, 14-IX-93 on Forsythia sp. (93T019). 

Ovatus crataegarius (Walker): ID, BONNER—Sandpoint, 10-IV-97 on Origanum sp. (97T003); WA, ADAMS— 
Othello, 19-VI-96 & 16-VII-96 on Mentha piperita (96B006, 96B010). 

Phorodon humuli (Schrank) [on Humulus lupulus|: WA, BENTON [*]—WSU-Prosser, 9-[X-95 (95K154). 

Rhopalosiphum insertum (Walker): WA, CHELAN—-WSU-Wenatchee, 30-V-96 on Pyrus malus (96RO117). 

Rhopalosiphum padi (L.) [on Triticum aestivum]: WA, BENTON—Rattlesnake Hills, 17-VII-95 (95G3577); 
WSU-Prosser, 30-IX-97 (97F457+, 97F463+); YAKIMA—Mabton Grade, 31-X-97 (97G517+). 

Sitobion avenae (F.): WA, BENTON—-WSU-Prosser, 11 & 18-VII-97 on Triticum aestivum (97F225, 97F243). 

Undetermined sp.: WA, BENTON—-WSU-Prosser, 29-X-95 on Senecio vulgaris (95G735). 


Aphidius nigripes Ashmead 
Macrosiphum euphorbiae (Thomas): WA, Kitrrras—Manastash Rd & FR112, 9-VU-96 on Agoseris elata 
(96G341) & Arnica cordifolia (96G332),; Lost Lk, 2-IX-95 on Aster sp. (95G528) & Castilleja sp. 
(95G527), & 11-IX-95 on Arabis sp. (95G568) & Monardella odoratissima (95G5697). 
Macrosiphum clydesmithi Robinson: WA, YAKIMA—Clear Lk, 15-VI-96 on Holodiscus discolor (96K081). 


Aphidius ohioensis Smith 

Macrosiphoniella ludovicianae (Oestlund) [on Artemisia ludoviciana]: CA. SHASTA [*]; WA, AsoTIN—Asotin 
Cr WLA, 5-V11—96, (96T138); YAKIMA—Wenas Lk, 17-VI-97 (97G090). 

Macrosiphoniella tanacetaria (Kaltenbach): WA, PIERCE—RNP, Tipsoo Lk, 5-IX-97 on Tanacetum vulgare 
(97G369). 

Macrosiphum creelii Davis: WA, Kittiras—Manastash Cr Rd, mi marker 8, 16-VIII-96 on Vicia sp. (96G492). 

Macrosiphum sp.?: ID, LATAH—Moscow Mtn, 6-VII-96 on Holodiscus discolor (961182). 

Macrosiphum valerianae (Clarke) [on Epilobium angustifolium|: WA, Kirrrras—Lost Lk, 24-VI-95 
(95G399++); S. Fork Manastash Cr, 24-VII-95 (95G381); YAKIMA—YIR, Signal Peak, 11-VII-95 
(95G322¢). 

Microsiphoniella artemisiae (Gillette): WA, AsoTiIn—Asotin Cr HMU, 5-VII-96 on Artemisia ludoviciana 
(96T141). 

Uroleucon adenocaulonae (Essig): WA, KLick1rAt—Trout Lk, 29-VI-97 on Adenocaulon bicolor (97G163). 

Uroleucon ur arnesense Robinson: MT, SANDERS—18 mi E of Thompson Pass, 29-VII-95 on Solidago, sp. 
(95G457). 


Uroleucon nr epilobii (Pergande): WA, SPOKANE—Mt Spokane SP, 27-VII-95 on Epilobium angustifolium 
(95T358). 


Uroleucon erigeronense (Thomas) [on Conyza canadensis unless otherwise indicated]: WA, CHELAN— 
Leavenworth, 16-IX-95 (95K141); Kitrrras—Lost Lk, 6-VIII-97 on Haplopappus hirtus (97G3407), & 


VOLUME 102, NUMBER 3 695 


11-IX-95 on Achillea millefolium (95G574); Manastash Rd & Rd 112, 11-IX-95 on Cirsium arvense 
(95G5647+); S. Fork Manastash Cr, 11-IX-95 on Achillea millefolium (95G563), & 24-VII-95 on Crepis sp. 
(95G3867); LINCOLN—Lk Roosevelt, Goldsmith CG, 31-VII-97 (97G302); PIERCE—RNP, 8-IX-95 on Aster 
sp. (95K106) & on Senecio sp. (95K1207), Tipsoo Lk, 5-IX-97 on Aster sp. (97G364+, 97G366), & 24- 
VII-97 on Compositae (97G252); YAKIMA—Grandview, 9-VII-95 (95G314); Pleasant Valley CG, 24-VII- 
97 on Crepis rucinata (97G258); Sunnyside, 23-X-95 & 13-X-96 (95G707, 96G645); YIR, nr Mt Adams 
Lk, 4-VHI-95 on Eriophyllum lanatum (95K092), Signal Peak, 11-VII-95 on Aster sp. (95K049+), Cedar 
Cr, 19-[X-95 on Achillea millefolium (95G620+), Klickitat Canyon (95G6287+), & White Deer Cr, 11-VII- 
95 on Crepis modocensis (95K0487*). 

Uroleucon nr erigeronense (Thomas): WA, LINCOLN—Lk Roosevelt, Goldsmith CG, 31-VII-97, on Aster sp. 
(97G307); SPOKANE—Peone, 20-VII-95 on Solidago sp. (95T278); YAKIMA—Mabton Grade, 23-V-96, on 
Balsamorhiza careyana (96G069); YIR, nr Mt Adams Lk, 4-VIII-95 on Hieracium sp. (95K0837). 

Uroleucon escalantii (Knowlton): WA, BENTON—ALE, 8-VI-95 on Chrysothamnus viscidiflorus (95G148). 

Uroleucon nr escalantii (Knowlton): WA, KLICKITAT—Box Springs, nr Bickleton, 29-VI-97 on Chrysothamnus 
sp. (97G178). ; 

Uroleucon gigantiphagum Moran: WA, YAKIMA—YIR, Signal Peak, 11-VII-95 on Solidago sp. (95G329). 

Uroleucon ivae Robinson [on Iva xanthifolia|: WA, BENTON [*]—ALE, Lower Snively Spring, 12-VIII-95 
(95K104); YAKIMA [*]—Grandview, 7,9 & 22-VII-95, & 30-VII-96, (95G309, 95G313, 95G366 & 
96G431); Harrah, 29-IX-97 (97G465): Selah, 10-VIII-95 (95G500); Wenas-Ellensburg Rd, Ellensburg Pass, 
7-VIU-96 (96G317). 

Uroleucon katonkae (Hottes) [on Aster sp.]: WA, YAKIMA—Wenas Lk, 17-VI-97 (97G099); YIR, Lost Springs, 
11-VII-95 (95K0497). 

Uroleucon olivei Moran: WA, BENTON—WSU-Prosser, 21-VII-94 on Iva xanthifolia (94R012). 

Uroleucon russellae (HRL) [on Anaphalis margaritacea unless otherwise indicated]: WA, LEwis—RNP., 8- 
IX-95 on Senecio sp. (95K120+); SKAMANIA—June Lk, 4-VIII-95 (95K101); SPOKANE—Mt Spokane SP, 
20-VH-95 & 24-VII-95 (95T294, 95T494); YAKIMA—YIR, Lost Springs (95G412) & Soda Springs 
(95G423), 25-VII-95. 

Uroleucon sonchi (L.): WA, YAKIMA—Grandview, 17-VII-95, 8-VII-96 on Sonchus oleraceus (95G348, 
96G326); Sunnyside, 23-X-95 on Sonchus sp. (95G703); Satus, 1-VI-96 on Sonchus sp. (96K064). 

Uroleucon suzannae ROBINSON: WA, Kittiras—Lost Lk, 6-VII-97 on Haplopappus hirtus (97G340*#). 

Uroleucon spp.: ID, BENEWAH—McCroskey SP, 31-VIII-95 on Adenocaulon bicolor (951527, 951536): WA, 
CHELAN—Chiwawa Canyon, 3-VIII-96 on Adenocaulon bicolor (96R044); Kirrrras—Lost Lk, 11-IX-95 
& 6-VII-97 on Aster sp. (95G579); Lost Lk Trailhead, 6-VIII-97 on Agoseris retrorsa (97G319); S. Fork 
Manastash Cr, 24-VII-95 on Hieracium sp. (95G389); LINCOLN—Lk Roosevelt, Goldsmith CG, 31-VI-97 
on Tragopogon dubius (97G303); PIERCE—RNP, Tipsoo Lk, 5-I[X-97 on Anaphalis margaritacea (97G367); 
SPOKANE—Mt Spokane SP, 24-VIII-95 on Centaurea pratensis (95T477); YAKIMA [on Iva xanthifolia *|— 
Hwy 410, Fife Peak Viewpoint, 24-VII-97 on Senecio serra (97G270+); Raven’s Roost, 28VII-97 on 
Hieracium gracile (97G2987); Sawmill Flats CG, 26-VII-97 on Aster sp. (97G287); Union Gap, 23-X-95 
on Centaurea maculosa (95G720); YIR, Big Spring, 4-VIII-95 on Agoseris elata (95KO090+), Lost Spring, 
25-VII-95 on Cichorium intybus (95G413), Signal Peak, 11-VH-95 on Eriophyllum lanatum (95KO055), 25- 
VII-95 on Achillea millefolium (95G410, 95G417); 19-[X-95 on Grindelia sp. (95G618), Signal Peak Guard 
Station, 19-IX-95 on Antennaria sp. (95G616), & Windy Point, 19-IX-95 on Aster sp. (95G629). 


Aphidius nr pisivorus Smith 
Acyrthosiphon pisum (Harris): WA, BENTON—-WSU-Prosser, 11-VII-97 on Melilotus officinalis (97F2187*). 


Aphidius polygonaphis (Fitch) 

Acyrthosiphon lactucae (Passerini) [on Lactuca serriola|: WA, WHITMAN—Kamiak Butte, 12-IX-95 (95T550); 
YAKIMA—Sunnyside, 28-VI-95 (95G257). 

Amphorophora parviflori Hill: WA, YAKIMA—N. Fork Tieton River & Rd 820, 27-VI-96 on Rubus parviflorus 
(96G304). 

Amphorophora sp.?: WA, LEwis—RNP, Stevens Canyon, 23-VII-96 on Arnica sp. (96G3597). 

Aphis ceanothi Clarke: WA, YAKIMA—Windy Point CG, 3-VI-95 on Ceanothus integerrimus (95G107). 

Aphis helianthi Monell: WA, YAKIMA—YIR, nr. Mt. Adams Lk., 25-VII-95 on Epilobium angustifolium 
(95G402+¢). 

Aphis nr oenotherae Oestlund: WA, GARFIELD—Stentz Springs, 14-IX-95 on Epilobium sp. (95T565). 

Brachycaudus cardui (L.): MT, SANDERS—Thompson Pass, 17-VI-95 on Senecio sp. (95G1917#). 

Capitophorus elaeagni (del Guercio): WA, KitrirAs—Frost Meadows, 11-[X-95 on Cirsium arvense 
(95G5837). 

Ericaphis wakibae (Hottes): WA, YAKIMA—Clear Lk, 15-VI-96 on native Rosa sp. (96K084, 96K0857). 


696 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Hyperomyzus nigricornis (Knowlton): WA, YAKIMA—Bird Cr Meadows, 26-VI-95 on Hieracium sp. 
(95A0167). 

Hyperomyzus sp. [nr nigricornis (Knowlton) & sandilandicus (Robinson)| WA, KitrrirAs—S. Fork Manastash 
Cr, 24-VII-95 on Hieracium albiflorum (95G388). 

Iillinoia nr azaleae (Mason): WA, Y AKIMA—Raven’s Roost, 28-VI-97 on Rhododendron albiflorum (97G297). 

Illinoia davidsoni (Mason): WA, LEwis—RNP, Stevens Canyon, 23-VII-96 on Arnica sp. (96G3597); 
YAKIMA—Bird Cr Meadows, 26-VI-95 on Rubus parviflorus (95A019). 

Illinoia sp. [on Vaccinium sp.]: ID, SHOSHONE—Thompson Pass, 29-VII-95 (95G472); WA, SKAMANIA— 
Keenes Horse Camp, 11-VII-95 (95G337). 

Macrosiphoniella ludovicianae (Oestlund) [on Artemisia ludoviciana|: WA, YAKIMA [on Artemisia 
ludoviciana* |—YIR, Bedground Springs, 16-V-96 (96G0537+). 

Macrosiphum clydesmithi (Robinson): WA, YAKIMA—YIR, Piscoe Meadows, 5-VIH-95 on Pteridium 
aquilinum (95G4337). 

Macrosiphum clydesmithi (Robinson): WA, SPOKANE—Mt Spokane SP, 20-VII-95 on unknown Polypodiaceae 
(95T298); YAKIMA—Clear Lk, 15-VI-96 on Holodiscus discolor (96K081). 

Macrosiphum euphorbiae (Thomas): MT, SANDERS—Thompson Pass, 17-VI-95 on Senecio sp. (95G1917); 
WA, AsoTiN—De Spain Spring, 11-VII-96 on Potentilla sp. (961250); 11-VII-96, on unknown plant 
(96T252); KittiraAs [on Rosa sp.*]|—Manastash Canyon, 2-1X-95 on Lactuca serriola (95G544); Lost Lk 
Trail, 6-VIII-97 on Luina nardosmia (97G327); WHITMAN—Pullman, 22-VII-95 on L. serriola (95T300) 
& 18-VI-96 on cultivated Rosa sp.(96T037); YAKIMA—Clear Lk, 15-VI—96 on native Rosa sp. (96K0857); 
Bird Cr Meadows, 26-VI-95 on Hieracium sp. (95SA016*). 

Macrosiphum nr euphorbiae (Thomas): WA, YAKIMA—YIR, Klickitat Canyon, 25-VII-95 on Cynoglossum 
grande (95G418); Mill Cr Guard Station, 6-VI-95 on Lomatium nudicaule (95G115). 

Macrosiphum pteridis (Wilson): OR, BENTON—McDonald Forest, 14-IV-94 on Holodiscus discolor, A. Jensen 
collector (94JOO1). 

Macrosiphum rosae (L.): OR, BENTON—Corvallis, 18-V-93 on unknown plant, A. Jensen collector (93J003). 

Macrosiphum valerianae (Clarke): WA, YAKIMA—YIR, nr. Mt. Adams Lk., 25-VIH-95 on Epilobium 
angustifolium (95G4027*). 

Macrosiphum sp.: ID, BENEWAH—McCroskey SP, 7-VI-96 on native Rosa sp. 96TO05); WA, GARFIELD— 
FR160 & FR40, 21-IX-95, on Streptopus sp. (9ST601); KittrrAs—Manastash Cr, mi marker 12, 6-VIII- 
97 on Holodiscus discolor (97G314); YAKIMA—YIR, Piscoe Meadows, 25-VII-95 on Pteridium aquilinum 
(95G4337+); Howard Lk, 19-IX-95 on P. aquilinum (95G640). 

Nasonovia crenicorna (Smith & Knowlton): WA, WHITMAN—Pullman, 27-V-97 on Geranium viscosissimum 
(97T035) 

Nasonovia houghtonensis (G&P): WA, WHATCOM—Birch Bay, 29-V-93 on Oemleria cerasiformis, A. Jensen 
collector (93J004). 

Sitobion sp.. CANADA, B.C.—Vancouver, Univ. of British Columbia, 29-V-93 on Dicentra formosa, A. 
Jensen collector (93J003). 

Uroleucon nr arnesense Robinson: WA, CHELAN—Leavenworth, Icicle Cr, 1-VIII-95, on Solidago canadensis 
(95G490). 

Uroleucon nr erigeronense (Thomas): WA, KittirAs—Manastash Rd & Rd112, 9-VII-96 on Hieracium 
albiflorum (96G340). 

Uroleucon ivae Robinson: WA, YAKIMA—Sunnyside, 13-VIH-94 & 11-VII-97, on Iva xanthifolia (94GO009, 
97G242). 

Wahlgreniella nervata (Gillette): ID, SHOSHONE—Thompson Pass, 29-VH-95 on Pachistima myrsinites 
(95G471). 


Aphidius nr polygonaphis (Fitch) 
Macrosiphum walkeri Robinson: OR, BENToN—Corvallis, 17-III-90 on Polypodium sp. (90J001). 


Aphidius pulcher Baker 
Acyrthosiphon pisum (Harris): WA, BENTON—-WSU-Prosser, 1-VII-96 on Melilotus officinalis (96G3117). 


Aphidius rhopalosiphi DeStefani 
Metopolophium dirhodum (Walker): WA, BENTON—-WSU-Prosser, 6-VIII-97 on Triticum aestivum (97F2267). 
Sitobion avenae (FE): WA, BENTON—-WSU-Prosser, 11, 18 & 25-VII-97 on Triticum aestivum (assorted 
collections between 97F204—97F277). 


Aphidius nr rosae Haliday 
Macrosiphum pallidum (Oestlund): WA, SPOKANE—nr Cheney, 23-V-95 on native Rosa sp. (95KO006). 


VOLUME 102, NUMBER 3 697 


Aphidius salicis Haliday 

Cavariella aegopodii (Scopoli): WA, BENTON—-WSU-Prosser, 21-VII-97 on Anethum graveolens (97G247}+); 
Kitritas—Reecer Cr, nr Ellensburg, 20-VI-95 on Angelica canbyi (95G203); Manastash Cr Rd, mi marker 
12, 25-VI-97, on Osmorhiza occidentalis (97G132); Lost Lk, 6-VIII-97 on Osmorhiza chilensis (97G331); 
Quilomene WLA, Crossover Rd & Colockum Ridge, 2-VH-97 on Osmorhiza purpurea (97G207); PEND 
OREILLE—Tiger Meadows, 9-VII-96 on unknown Umbelliferae (96T219+); YAkIMA—Hwy 410, Fife Peak 
Viewpoint, 24-VH-97 on Osmorhiza chilensis (97G271+); YIR, Signal Peak, 19-Ix-95 on Salix sp. 
(95G623). 

Cavariella pastinacae (L.) [on Heracleum lanatum unless otherwise indicated]: WA, AsoTIN—De Spain 
Springs, 5-VII-96 & 25-VII-96 (96T 156, 96T350); ISLAND—N. Whidby Island, 5-VI-96 (96G171); KING— 
Tinkham exit, 4-VI-96 (96G137); KirrirAs—Reecer Cr, nr Ellensburg, & S. Fork Manastash Cr, & 
Manastash Rd, 20-VI-95, 24-VII-95, & 25-VII-96 (95G217, 95G393, 96G417); Taneum Lk, 11-IX-97 on 
Angelica sp. (97G413); PIERCE—RNP, Ipsut Pass, 25-[X-97 (97G447); Mowich Lk, 25-IX-97 (97G450); 
YAKIMA—Yakima, 28-VI-95 on Heracleum lanatum (95G269+); YIR, Klickitat Canyon, 23-VI-95 
(95G235), & Signal Peak, 11-VII-95 (95G335). 


Aphidius segmentatus Pike and Stary, new species 
(Figs. 1-9) 


Diagnosis.—The new species is distinguished by several characters, but especially a 
short antenna (14—15 segments, noticeably widening to apex), and short distal abscissa 
of Rl (= metacarpus). In Smith (1944), the new species keys to Aphidius ribis Haliday, 
but the latter’s antenna is longer, not remarkably thickened, and the metacarpus is long. 
Similar differences also distinguish the new species from a European species, A. sonchi 
Marshall that attacks Hyperomyzus aphids (Stary 1963, 1966). 

Etymology.—The name of the new species is derived from the somewhat unusual 
segmentation of the flagellum. 

Description.—Female. Head: Eye length 3X malar space. Tentorio-ocular line % 
intertentorial line. Antenna 14—15 (prevalently 15) segmented, short, as long as head, 
mesosoma and metasomal tergum | together, thickened to apex, segmentation of flagellum 
rather distinct (Fig. 7). Flagellomeres with semierect setae (Figs. 4—6). Flagellomere 1 
[F,] (Fig. 4) length 2 width, with O—1 longitudinal placodes, setae about 74 segment 
diameter. F, similar to F,, with 2—3 placodes. Middle F (Fig. 5) length about 1.3 width, 
distinctly wider than F,. Preapical F (Fig. 6) length about 2 width, wider than middle 
E Mesosoma: Mesonotum glabrous, with rows of sparse setae indicating the effaced 
notauli on the disc. Propodeum (Fig. 1) with rather small areola, carinae sometimes 
indistinct to slightly indicated in upper portion. Forewing: (Fig. 8): Stigma length 3X 
width; distal abscissa of R1 (metacarpus) equal or subequal to stigma width. Vein r slightly 
longer than stigma width, and twice RS length; r-m equal to RS in length. Marginal setae 
3X length of surface setae. Metasoma: Metasomal tergum 1 (Fig. 9) length 2.5 width 
at spiracles; surface slightly rugose in anterior portion; anterolateral area costulate. 
Genitalia (Fig. 2): Ovipositor sheath (Fig. 3) apex blunt. Coloration: Head brown; face, 
gena, and mouthparts yellow, except brown apices of mandibles. Antenna brown; a ring 
between pedicel and flagellum sometimes yellow. Mesosoma brown, propleuron yellow; 
mesopleuron sometimes with yellow markings in lower portion. Tegula yellow. Wings 
hyaline, venation brown. Legs light brown; coxa, trochanter and base of tibia yellow; a 
dark tinge increasing from fore to hind legs, the latter the darkest. Metasoma: Tergum 1 
and apical half of metasoma yellow, terga 2 and 3 and some times 4, with brown banding. 
Ovipositor sheath brown. Length of body: about 2.2—2.7 mm. 

Male.—Antenna 17—18 segmented. Coloration: head brown; mouthparts mainly (apices 
of mandibles brown, palpi light brown). Mesosoma brown. Legs brown, base of tibia 
yellow to light brown. Metasoma brown, tergum 1 yellow to light brown. 


698 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-9. Aphidius segmentatus, 2 {illustrations not to equal scale]. 1, Propodeum. 2, Genitalia. 3, Ovi- 
positor sheath. 4, Flagellomere 1 [F,]. 5, Middle flagellomere. 6, Preapical flagellomere. 7, Antenna [flagellum]. 
8, Fore wing, in part. 9, Metasomal tergum 1. Abbreviations: R1, distal abscissa of postmarginal vein [= 
metacarpus]; RS, r, and r-m, wing veins. 


VOLUME 102, NUMBER 3 699 


Holotype.—?, USA, WA, YAKIMA Co., Yakama Indian Reservation, 25-VII-1995, G. 
Graf collector, (collection code, 95G406). Host aphid: Hyperomyzus nigricornis 
(Knowlton) on Hieracium cynoglossoides. Deposited in USNM. 

Paratypes.—16 specimens, collection data same as holotype, deposited in part in 
collections of WSU-Prosser (10 2, 4 d) and P. Stary, Ceské Budejovice, Czech Republic 
(1 2,1); 11 specimens, USA, WA, Kittitas Co., L.T. Murray WLA, S. Fork Manastash 
Creek, 24-VII-1996, on Hyperomyzus sp. (nr nigricornus or sandilandicus) on Hieracium 
albiforum (96G388), deposited in part in collections of WSU-Prosser (6 ¢) and P. Stary, 
Ceské Budejovice, Czech Republic (3 2, 2 ¢). 

Other material reared from.— 

Hyperomyzus lactucae (L.): WA, WHItMAN—Pullman, 12-IX-95, on Lactuca serriola 
G5m559). 

Hyperomyzus nigricornis (Knowlton) [on Hieracium sp. unless otherwise indicated]: 
ID, LATAH—Moscow Mtn, 3-VII-96 on Compositae (96T399); WA, GARFIELD—FR40 nr 
Teal Springs, 1-VIII-95 on Senecio sp. (95T378+); Pataha Cr Rd, 25-VII-95 (9513234); 
Wickiup CG, 29-VIII-95 (95T508); KittiraAs—Lost Lk, 6-VIII-97 (97G317+, 97G344); 
KLICKITAT—Cleveland, 7-VII-97 (97G229); PIERCE—RNP, Paradise, 8-[X-95 on Senecio 
sp. (95K119), & Tipsoo Lk, 5-IX-97 (97G359); SPOKANE—Mt Spokane SP, 24-VIII-95 
on Compositae (95T486); YAKIMA—Little Naches River, 5 mi W of Timothy Meadows, 
9-IX-97 (97G388, 1 specimen); Hwy 410, Ravens’s Roost, 28-VII-97 (97G2987+); Pleasant 
Valley CG, 24-VII-97 (97G2597); Sawmill Flats, 28-VII-97 (97G2887); Wenas Lk, 17- 
VI-97 on Compositae (97G087); R, Signal Peak, 11-VII-95 (95K056). 

Hyperomyzus sp. [nigricornis (Knowlton) or sandilandicus (Robinson)]: WA, 
KitTTITAs—Manastash Rd & FR112, 11-IX-95 on Hieracium sp. (95G5597); L. T. Murray 
Wildlife Reserve, South Fork Manastash Cr, 24-VII-95 on Hieracium albiflorum 
(95G388). 

Uroleucon erigeronense (Thomas): WA, KittirAs—Manastash Ridge, nr Lost Lk, 2- 
IX-95 on Achillea millefolium (95G533¢£). 


Aphidius smithi Sharma & Subba Rao 
Acyrthosiphon pisum (Harris): WA, BENTON [on Medicago sativay;*]. 


Aphidius spp. 

Acyrthosiphon lactucae (Passerini) [on Lactuca serriola}: 1D, BENEWAH—McCroskey SP, 31-VIII-95 
(95T534); WA, Asotin—Asotin, 10-VII-95 (95T419); CHELAN—Chelan SP, 11-IX-97 (97K038); 
SPOKANE—nr Peone, 24-VIII-95 (95T470). 

Acyrthosiphon macrosiphum (Wilson) [on Amelanchier alnifolia]: MT, LINCOLN—Libby, 6-VIH-96 (96G322); 
WA, PEND OREILLE—Le Clere WLA, Panhandle CG, 3-VIII-95 (95T402). 

Acyrthosiphon pisum (Harris): WA, BENTON [on Medicago sativa*]; Kittiras [on Rhododendron albiflorum*}). 

Amphorophora geranii G&P: WA, SPOKANE—Turnbull National WLA, 25-V-95 on Geranium sp. (95T0307). 

Amphorophora nr rubitox’ 1 Knowlton: WA, Is_AND—South Whidbey Island, 4-VI-96 on Rubus sp. (96G163). 

Aphis armoraciae Cowen: WA, Kittitas [on Mertensia paniculata*}. 

Aphis craccivora Koch: WA, WHITMAN—Pullman, 5-VII-97 on Trifolium sp. (96T170). 

Aphis fabae Scopoli: WA, AsoTiN—Field Springs SP, 18-VII-95 on Cirsium arvense (951272). 

Aphis helianthi Mo ell: WA, BENTON—-WSU-Prosser, 21-VU-97 on Anethum graveolens (97G247*). 

Aphis holodisci Robinson: WA, AsoTIN—nr FR40 & FR44, 26-VI-96 on Holodiscus discolor (96T052). 

Aphis nigratibialis Robinson: WA, YAKIMA—Nile, 28-VII-97 on Cornus stolonifera (97G280). 

Aphis varians Patch: WA, PIERCE [on Epilobium angustifolium* }. 

Aphis (Zyxaphis) sp.: WA, KLick1rat—Frost Meadows, 11-[X-97 on Artemisia tridentata (97G435). 

Aspidophorodon longicauda (Richards): WA, SKAMANIA—South Prairie, 8-IX-94 on Spiraea douglasii 
(94R192). 

Brachycaudus helichrysi (Kaltenbach): WA, SPOKANE—Plaza, 16-VII-96 on unknown plant (96T278). 

Brachycaudus tragopogonis (Kaltenbach): WA, BENTON—WSU-Prosser, 25-VH-97 on Tragopogon dubius 
(97F 266). 


700 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Capitophorus elaeagni (del Guercio): WA, YAKIMA—Rimrock Lk, 14-[X-95 on Cirsium vulgare (95G589). 

Carolinaia howardii (Wilson): WA, WHITMAN—Pullman, 1-VII-96 on unknown plant (96T123). 

Chaetosiphon fragaefolii (Cockerell): WA, PEND OREILLE—Le Clerc WLA, 3-VIII-95 on native Rosa sp. 
(951398); STEVENS—Little Pend Oreille National WLA, 9-VII-96 on native Rosa sp. (96T2097). 

Chaetosiphon nr hottesi Stroyan: WA, SPOKANE—Mt Spokane SP, 24-VIII-95 on native Rosa sp. (9ST4917£). 

Chaetosiphon tetrarhodum (Walker): WA, SPOKANE—Mt Spokane SP, 24-VIII-95 on native Rosa sp. 
(95T49174). 

Diuraphis noxia (Kurdjumov) [on Triticum aestivum]: WA, AsoTiIN—Field Springs SP, 20-VI-95 (95T158); 
BENTON [*]. 

Ericaphis gentneri (Mason) [on Crataegus sp.]: WA, CHELAN—Stevens Pass, 16-I[X-95 on Spiraea densiflora 
(95K 138); Kirriras—Coleman Canyon, 2-VII-97 (97G214); SPOKANE—Chapman Lk, 16-VII-96 (96T282). 

Ericaphis nr gentneri (Mason): WA, WHITMAN—Pullman, 6-VIII-96 on Crataegus (96T409). 

Ericaphis scammelli (Mason): WA, STEVENS—Little Pend Oreille National WLA, 9-VII-96 on native Rosa 
sp. (96T2097). 

Ericaphis wakibae (Hottes): WA, YAKIMA—N Fork Tieton River, 15-VI-96 on native Rosa sp. (96K090*). 

Euceraphis gillettei Davidson: WA, KitrirAs—L.T. Murray WLA, S Fork Manastash Cr, 24-VI-95 on Alnus 
sp. (95G3947+). 

Hyperomyzus nigricornis (Knowlton): WA, Kittiras—Lost Lk, 6-VIII-97 on Hieracium sp. (97G344). 

Hyperomyzus sp.: WA, BENTON [on Sonchus oleraceus*]. 

Illinoia crystleae bartholomewi (Essig) [det. by A. Jensen]: WA, Kittiras—Lost Lk, on Lonicera involucrata 
(97G336). 

Illinoia nr crystleae (Smith & Knowlton): WA, Kitriras—Lost Lk, 7-VII-94 on Lonicera involucrata 
(94K028). 

Illinoia davidsoni (Mason): WA, GARFIELD—Alder Thicket, 22-VIII-95 on Arnica latifolia (95T450). 

Illinoia rhododendri (Wilson): WA, Kittitas [on Rhododendron albiflorum* |. 

Illinoia spp.: WA, GARFIELD—Stentz Springs, 1-VIII-95 on Aquilegia formosa (9513627); SkAGiIt—La Conner, 
6-VI-96 on Rhododendron sp. (96G193); SKAMANIA [on Vaccinium sp.*]. 

Liosomaphis berberidis (Kaltenbach): WA, SkAGiIr—La Conner, 5-VI-96 on Berberis sp. (96G192). 

Macrosiphoniella ludovicianae (Oestlund): WA, Asotin—Anatone, 18-VIH-95 on Artemisia ludoviciana 
(Osik257)): 

Macrosiphum californicum (Clarke) [on Salix sp.]: WA, Kittitas [+*]; YAKIMA—YIR, Lost Springs, 11-VII- 
95 (95KOS52). 

Macrosiphum clydesmithi (Robinson): WA, CHELAN—Chiwawa Canyon, 5-IX-95 on Pteridium aquilinum 
(95RO0267). 

Macrosiphum euphorbiae (Thomas): ID, LATAH—Moscow Mtn, 6-VII-96 on unknown plant (961187); 6-VII- 
96 on Compositae (96T185); MT, SANDERS—Thompson Pass, 17-VI-95 on Hieracium sp. (95G185), & 
Arabis sp. (95G190+); WA, GARFIELD—Stentz Springs, 1-VII-95 on Arnica latifolia (95T370); Spruce 
Springs, 22-VIII-9, on Ranunculus sp. (95T459); Kittiras—Manastash Rd 25-VII-96 on Ranunculus sp. 
(96G416); Taneum Lk, 11-IX-97 on Silene sp. (97G415); Lost Lk, 6-VIII-97 on Angelica canbyi 
(97G330+); KLickirat—Cleveland Pk, 7-VII-97 on Lupinus sp. (97G230); SPOKANE—Mt Spokane SP, 29- 
VI-95 on Ranunculus sp. (95T190); WHITMAN—Pullman, 5-VII-96 on cultivated Rosa sp. (9611727); 
YAKIMA—N. Fork Tieton River, 27-VI-96 on Hieracium scouleri (96G297); Buckhorn Meadows, 14-IX- 
95 on Trautvetteria caroliniensis (95G597). 

Macrosiphum pallidum (Oestlund): MT, SANDERS—Thompson Pass, 17-VI-95 on Arabis sp. (95G1907#). 

Macrosiphum rhamni (Clarke) [det. by A. Jensen]: WA, Kinc—Stampede Pass, 8-VIII-96, on Pteridium 
aquilinum (96G441); KLickiIrat—Trout Lk, 29-VI-97 on Pteridium aquilinum (97G169). 

Macrosiphum rosae (L.): WHITMAN—Pullman, 5-VII-96 on cultivated Rosa sp. (9611727). 

Macrosiphum tuberculaceps (Essig): WA, YAKIMA—Hell Crossing CG, 11-[X-96 on Achlys triphylla 
(96G517); N. Fork Tieton River, 15-VI-96 on A. triphylla (96K096). 

Macrosiphum sp.: WA, Ktinc—Kirtiras—Waptus Lk, 27-VII-95 on Spiraea douglasii menziesii (95KO75),; 
Manastash Cr, mi marker 12, 25-VI-97 on Holodiscus discolor (97G128); SPOKANE—Turnbull National 
WLA, 25-V-95 on Geranium sp. (95T0307); YAKIMA—Nile, 28-VII-97 on Clematis ligusticifolia (97G278). 

Metopolophium dirhodum (Walker): WA, Kittitas [on Rhododendron albiflorum*]|; SNoHOMISsH—Monroe, 5- 
VI-96 on Phalaris arundinaceae (96G202+); YAKIMA—N Fork Tieton River, 15-VI-96 on native Rosa sp. 
(96K0907). 

Myzus cerasi (FE): WA, SKAGIT—Campbell Lk, 5-VI-96 on Prunus emarginata (96G170). 

Nasonovia aquilegiae (Essig): WA, GARFIELD—Stentz Springs, 1-VIII-95 on Aquilegia formosa (9513627). 

Nasonovia nr aquilegiae (Essig): WA, SPOKANE—Manito Pk, 11-VII-96 on Aquilegia sp. (9612627). 


VOLUME 102, NUMBER 3 701 


12 


Figs. 10-12. Betuloxys alnicolus, 2 [illustrations not to equal scale]. 10, Metasomal tergum 1. 11—12, Prong. 


Nasonovia polemonii (G&P): WA, Kittiras—Buck Meadows, 9-VII-96 on Polemonium pulcherrimum 
(96G347). 

Nasonovia sp.: MT SANDERS [on Castilleja sp.*]. 

Oestlundiella flava (Davidson): WA, KittrrAs—L.T. Murray WLA, S Fork Manastash Cr, 24-VII-95 on Alnus 
sp. (95G39474). 

Ovatus crataegarius (Walker): ID, BONNER—Sandpoint, 10-IV-97 on Origanum sp. (97T003); WA, ADAMS— 
Othello, 1-VII-96 on Mentha piperita (96B003). 

Prociphilus sp.: WA, YAKIMA—Clear Lk, 15-VI-96 on Lonicera sp. (96K083). 

Rhopalosiphum padi (L.): WA, SNOHOMISsH—Monroe, 5-VI-96 on Phalaris arundinaceae (96G2021). 

Sitobion avenae (F.): WA, BENTON—-WSU-Prosser, 18-VII-97 on Setaria lutescens (97G243). 

Sitobion fragariae (Walker): WA, Is_LAND—Whidbey Island, 5-VI-96 on Dactylis glomerata (96G177). 

Uroleucon erigeronense (Thomas): WA, YAKIMA—Buckhorn Meadows, 14-IX-95 on Achillea millefolium 
(95G598). 

Uroleucon ivae Robinson [on Iva xanthifolia}: WA, BENTON [*]; YAKIMA [*]—Grandview, 21-VIII-95 
(95G503). 

Uroleucon russellae (HRL): WA, SKAMANIA—Council Lk, 4-VIII-95 on Anaphalis margaritacea (95K095). 

Uroleucon sonchi (L.): WA, BENTON [on Sonchus oleraceus**}. 

Uroleucon sp.: WA, PEND OREILLE—Sullivan Lk, 3-VIII-95 on Adenocaulon bicolor (957391); Tiger 
Meadows, 9-VII-96 on Senecio sp. (96T220). 

Utamphorophora humboldti (Essig): OR. MULTNOMAH [on Physocarpus sp.*]. 

Undetermined spp.: WA, AsoTIN [on Chrysothamnus nauseosus*]; DOUGLAS—Waterville, 13-VII-95 on 
Triticum aestivum (95B013, 95B014); KittrrrAs—Quartz Mtn 25-VII-96 on Rubus lasiococcus (96G407): 
Taneum Lk, 11-IX-97 on Pedicularis bracteosa (97G425); LEwIs—RNP., Louise Lk, 1-VII-95 on Luetkea 
pectinata (95K042),; SPOKANE—Mt Spokane SP, 22-VII-96 on Vaccinium sp. (96T492); YAKIMA—Clear 
Lk, 15-VI-96 on Holodiscus discolor (96K080); YIR, Mabton Rd, 16-V-96 on Grindelia sp. (96G051). 


Genus BETULOXYS Mackauer 
Betuloxys alnicolus Pike & Stary, new species 
(Figs. 10—12) 


Diagnosis.—The new species is easily distinguished from B. compressicornis (Ruthe) 
by its distinctly smaller size, generally brown coloration, and characters on prongs (distal 
portion less differentiated and bearing three lanceolate setae). The new species was col- 
lected near colonizing aphids [Boernerina variabilis (Richards)] on Alnus in Washington. 
By comparison, B. compressicornis is generally yellow (exception, flagellum mostly 
brown, and brown areas on mesonotum, scutellum, apices of legs and prongs), distal 
portion of prong with five lanceolate setae, known as a parasitoid of Euceraphis puncti- 
pennis (Zett.) on Betula, from E. Canada and Europe. 

Etymology.—The name of the new species is derived from its association with Alnus. 

Description.—Female. Head: Eyes medium sized; antenna 12-segmented, filiform, 
reaching to half of metasoma. Mesosoma: Mesonotum smooth, with sparse setae, notauli 
indicated anteriorly. Propodeum feebly granulate rugose, with sparse setae. Forewing: 


702 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Stigma slightly more than 3X as long as wide (length/width [L/W] = 3.2); distal abscissa 
of R1 (= metacarpus) 2 width of stigma, about % shorter than stigma length; RS vein 
long, 1.2 stigma length. Metasoma: Metasomal tergum 1 (Fig. 10) smooth, length 1.6 
width, spiracular tubercles distinct, prominent laterally, situated at end of first third of 
tergite. Genitalia (Figs. 11-12): Prongs arcuate, apical portion somewhat dilated with 
three lanceolate, perpendicular setae on dorsal side. Coloration: Head and mesosoma dark 
brown. Mouthparts light brown. Antenna largely brown, except scape, pedicel, and F, 
yellow, F, yellow brown. Wings subhyaline, venation light brown. Legs brown. Metasoma 
brown, tergum 1 somewhat darker at base. Ovipositor sheaths and prongs brown, con- 
colorous with metasoma. Length of body: about 1.8 mm. 

Male.—Unknown. 

Holotype.—?, USA, WA, YAKIMA Co., N. Fork of Tieton River, 15-VI-1996, K. S. 
Pike collector (collection code, 96K089). Host aphid uncertain, collected near colonizing 
aphids, Boernerina variabilis Richards on Alnus. Deposited in USNM. 


Genus BINODOXYS Mackauer 


Binodoxys carolinensis (Smith) 

Aphis ceanothi Clarke: WA, KitrirAs—S. Fork Manastash Cr, 25-IV-97 on Ceanothus velutinus (97G137). 

Aphis helianthi Monell: WA, CHELAN—Johnny Cr, 1-VIII-95 on Epilobium angustifolium (95G4827); PEND 
OREILLE—Le Clerc WLA, 3-VIII-95 on Cornus stolonifera (95T399*+), YAKIMA—YIR, Signal Peak, 11- 
VII-95 on Lomatium sp. (95G3347). 

Aphis nigratibialis Robinson [on Cornus stolonifera]: WA, KLickirat—Trout Lk, 28-VI-97 (97G156); 
YAKIMA—Nile, 28-VII-97 (97G280); YIR, Miller Point, 11-VII-95 (95K062). 

Aphis nr oenotherae Oestlund: WA, YAKIMA—Pleasant Valley CG, 24-VH-97 on Epilobium minutum 
(97G2677). 

Aphis salicariae Koch: WA, PEND OREILLE—Le Clerc WLA, 3VIII—95 on Cornus stolonifera (95T399¥%). 

Aphis varians Patch [on Epilobium angustifolium]: 1D, SHOSHONE—Thompson Pass, 29-VII-95 (95G4687); 
MT, SANDERS [*]. 

Aphis nr varians Patch: WA, LEwis—RNP, Stevens Canyon, 23-VII-96 on Epilobium angustifolium (96G380). 

Aphis sp. n.: WA, YAKIMA—YIR, Miller Point, 11-VII-95 on Ceanothus sp. (95K061). 

Aphis sp.: WA, YAKIMA—Pleasant Valley CG, 24-VII-97 on Epilobium minutum (97G2677). 

Cavariella nr aegopodii (Scopoli): WA, KirtrrAs—nr Ellensburg, Reecer Cr, 20-VI-95 on Salix sp. (95G207#). 


Binodoxys clydesmithi Pike & Stary 
Aphis canae Williams: WA, FRANKLIN—Kahlotus Hwy, 11-VI-96 on Artemisia tridentata (96G242*%). 
Artemisaphis artemisicola (Williams) [on Artemisia tridentata]: WA, FRANKLIN—Kahlotus Hwy, 28-V-96 & 
11-VI-96 (96G096, 96G2427+#). 
Obtusicauda coweni (Hunter): WA, KITTITAS [on Artemisia tridentata* |. 


Binodoxys conei Pike & Stary 
Phorodon humuli (Schrank): WA, BENTON [on Humulus lupulus*]. 


Binodoxys coruscanigrans (Gahan) 
Obtusicauda coweni (Hunter): WA, YAKIMA [on Artemisia tridentata*|—Wenas Lk, 30-V-96, (96G115). 


Binodoxys grafi Pike & Stary 

Aphis fabae Scopoli: WA, Yakima—American River, 5-IX-97 on Epilobium angustifolium (97G377}). 

Aphis helianthi Monell [on Epilobium angustifolium]: WA, CHELAN—Stevens Pass, 16-IX-95 (95K1347); 
Kittitas [*]—Lost Lk, 24-VII-95 (95G399+); SPOKANE—Mt Spokane SP, 27-VII-95 on unknown 
Umbelliferae (95T3357). 

Aphis holodisci Robinson: WA, KitrrrAS—Manastash Cr, mi marker 12, 25-VI-97 on Holodiscus discolor 
(97G133). 

Aphis salicariae Koch: ID, KooTENAI—Rathdrum, 29-VII-95 on Epilobium angustifolium (95G443*). 

Aphis nr salicariae Koch: WA, CHELAN—Swauk Pass, 1-VIII-95 on Epilobium angustifolium (95G4927). 

Aphis nr triglochinis Theobald: WA, SPOKANE—Mt Spokane SP, 29-VI-95 on native Ribes sp. (95T189). 

Aphis varians Patch [on Epilobium angustifolium]: ID, KOOTENAI—Rathdrum, 29-VIH-95 (95G4437); WA, 


VOLUME 102, NUMBER 3 703 


CHELAN—Stevens Pass, 16-IX-95 (95K134+); Kittiras—Lost Lk, 24-VII-95 (95G399+); Stampede Pass, 
8-VIII-96 (96K 139). 
Aphis nr varians Patch: WA, Kittiras—Lost Lk, 24-VII-95 on Epilobium angustifolium (95G399+). 


Binodoxys rhagii (Ashmead) 
Uroleucon russellae (HRL) [on Anaphalis margaritacea |: WA, CHELAN—Chatter Cr, 1-VIII-95 (95G484); 
YAKIMA—YIR, Delaney Springs (95G412) & Panther Butte (95G423), 25-VII-95. 


Binodoxys spp. 
Aphis salicariae Koch: WA, YAKIMA [on Cornus stolonifera* ]. 
Chaetosiphon fragaefolii (Cockerell): WA, PEND OrEILLE—Le Clerc WLA, 3-VIII-95 on native Rosa sp. 
(95T398). 
Illinoia rhododendri (Wilson): WA, Kittitas [on Rhododendri albiflorum* |]. 


Genus DIAERETELLUS Stary 


Diaeretellus palustris Stary 
Rhopalosiphum nymphaeae (L.): WA, YAKIMA—Outlook, 23-X-95 on Typha latifolia (95G709). 


Genus DIAERETIELLA Stary 


Diaeretiella rapae (M Intosh) 

Acyrthosiphon lactucae (Passerini) [on Lactuca serriola]: WA, AsoTin—Asotin, 10-VIII-95 (95T419); S. Fork 
Asotin Cr, 22-VIII-95 (95T468); BENTON—WSU-Prosser, 17-VII-95 (95G353): Whitstran, 18-IX-97 
(97G439); WHITMAN—Pullman, 16 & 30-VII-96, 1,6 & 19-VIII-96, 9 & 12-IX-96 (961304, 96T362, 
96T382, 961407, 96T408, 96T468, 96T528, 961532). 

Acyrthosiphon pisum (Harris): MT, RAvALLI—Stevensville, 25-V-96 on Melilotus sp. (96G083); WA, 
BENTON—WSU-Prosser, 18 & 25-VII-97, on Melilotus officinalis (97F254+, 97F274+). 

Aphis craccivora Koch: WA, WHITMAN—Pullman, 25-VII-95 on Capsella bursa-pastoris (951331). 

Aphis fabae Scopoli: ID, LaTAH—Moscow Mtn, Paradise Cr, 25-IX-96 on Cirsium arvense (96T542). 

Aphis nr fabae Scopoli: WA, PIERCE—RNP, Paradise, 8-I[X-95 on Spiranthes romanzoffiana (95K118). 

Aphis helianthi Monell: WA, GARFIELD—Stentz Springs, 1-VIII-95 on Ranunculus sp. (9513637). 

Brachycaudus cardui (L.): WA, AsoTin—De Spain Springs, 5-[X-96 on Onopordum acanthium (96T518). 

Brachycaudus helichrysi (Kaltenbach): ID, BENEWAH—McCCroskey SP, 31-VIII-95 on Conyza canadensis 
(95T516+); WA, AsoTiIn—Anatone, 28-VI-96 on Amsinckia sp. (96T089); Wickiup CG, 29-VIII-95 on 
Sonchus sp. (95T505); BENTON—-WSU-Prosser, 1-X-97 on S. tuberosum (97G478); WHITMAN—Pullman, 
28-VI-96 on Anthemis arvensis (96T162); 16-VII-96 on Anthemis cotula (96T298). 

Brachycaudus tragopogonis (Kaltenbach) [on Tragopogon dubius]: WA, BENTON—WSU-Prosser, 3, 18, 25- 
VII-97 (97F194, 97F242, 97F266); KLickirat—Bickleton, 8-X-96 (96G628): YAKIMA—Grandview, 24-V- 
96, 25-VI-96 (96G080, 96G273). 

Brachycorynella asparagi (Mordvilko) [on Asparagus officinalis]: WA, BENTON [*]|—WSU-Prosser, 6 & 25- 
IX-95 (95K144, 95G548+, 95G649); YAkimA—Harrah, 29-IX-97 (97G462, 97G468, 97G469). 

Braggia sp.[det. by G. Remaudiére]: WA, YAKIma [on Eriogonum elatum* |. 

Brevicoryne brassicae (L.): WA, BENTON [on Brassica napus, B. oleracea & Descurainia sophia* |—WSU- 
Prosser, 6-VII-95, 24-VIII-95, 5-VIII-96, 30-IX-96, 17-I-97, 23-V-97, & 1-X-97 on B. napus (95A024, 
95G508, 96G438, 96G622, 97G004, 97G039, 97G475): 9-IX-95 on B. oleracea capitata (95K155); 11- 
VII-97 on Sisymbrium altissimum (97G241); 1-X-97, on Malva neglecta (97G471+): West Richland 12- 
VII-95 on B. oleracea acephala (95K074); CHELAN—Chelan SP. 11-[X-97 on Erysimum sp. (97K045); 
CLARK—WSU-Vancouver, 21-IX-95 on B. oleracea capitata (95K179); SkAGIr—Mt Vernon, 16-X-95 on 
B. sp. (95K190); PieRcE—Graham, 8-IX-96 on Cruciferae (961525); WALLA WALLA—Hwy 124, 10 mi E 
of Burbank, 28-V-96 on B. napus (96G099); 28-V-96 on Sisymbrium altissimum (96G098); WHITMAN [on 
B. napus*|—Pullman, 2-VIII-96 on B. napus (967390); YAKIMA— nr Bickleton 7-VII-97 on B. napus 
(97G235); on Sisymbrium altissimum (97G237). 

Diuraphis noxia (Kurdjumov) [on Hordeum vulgare unless otherwise indicated]: OR, UMATILLA—Hermiston, 
17-VI-90 on unknown plant (90J003); WA, Asotin—Columbia Center, Pataha Cr Rd, 10-VIII-95 on 
Agropyron cristatum (957431); Anatone, 24-VII-96, 1,8 & 15-VIII-96 (96T596 to 96T600): Anatone, 22 
& 31-VII-97 (97J0O11 to 97J025); BENTON [*]—WSU-Prosser, 28-V-96 & 24-X-96 (96G087 to 96G090, 
96G654); [KLICKITAT, on Aegilops cylindrica* |—Cemetery Rd nr Bickleton, 7-VII-97 (97G226+): WALLA 
WALLA—Valley Grove, 11-VI-96 (96G240); Waitsburg, 11-VII-96 (96J056, W. Turner & J. Rumph 
collectors); WHITMAN—Pullman, 9-VII-96 (96J053 to 96J055, W. Turner & J. Rumph collectors); YAKIMA 


704 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


[*]; [Following on Triticum aestivum unless otherwise indicated ] ADAMs [*]; ASOTIN [*|—Couse Cr Rd, 
10-VIII-95 (95T421); Field Springs SP, 10-VIII-95 (95T427); Asotin WLA, 15 & 29-VIII-96, 24-IX-96, 
3-X-96 (961455, 96T457, 96T503, 961555, 961569); De Spain Spring, 5-IX-96, 3-X-96 (96T516, 
96T571); BENTON [*]—WSU-Prosser, 15-V-95, 24-V-96, 3-VI-96, 24, 28-VII-97 (95G065, 96G081, 
96G132, 97G274, 97G301); 29-IV-97 thru 24-IX-97, (97FO01—97F004, 97F042—97F455); 1-IV-97, 6-VI- 
97 (97G005, 97G040, 97G041); 3 mi S of Prosser, 6-VI-97 (97G045); Sellards & Travis Rds, 25-VI-95 
(95G252); Rotha Rd 17-VII-97 (97G245); DouGLAas—Waterville, 12-VIII-95 (95B019); FRANKLIN [*]— Star 
School, 6-VII-95 (95G287); Kahlotus, 6-VII-95 (95G291); GARFIELD—Columbia Center, 10-VIII-95, 7-IX- 
95 (957435, 951538, 951539); Spruce Springs, 14-[X-95 (951574); KuickiTaT [*]—Bickleton, 31-X-95, 
7-VII-97 (95G743, 97G223): nr Bickleton, 18-VI-97 & 7 & 29-VII-97, 6-VIII-97 (97G119, 97G238, 
97G300, 97G354); Brannon Rd nr Cleveland, 7-VII-97 (97G234); Cleveland, 18-VI-97, 27-VI-97, 7-VII- 
97 (97G114 to 97G117, 97G145, 97G232); SPOKANE—Mt Spokane SP, 22-VII-95 (95T495); WHITMAN 
[*]|—Hole in the Ground, 8-VI-95 (95T085); Dusty, 9-VII-96 on Triticum sp. (96J065, W. Turner & J. 
Rumph collectors); YAKIMA [*]—nr Bickleton, 23-V-96, 17-VI-96 (96G073, 96G250, 96G251, 96G252); 
Sunnyside, 24-VH-97 (97G273). 

Dysaphis plantaginea (Passerini): WA, BENTON—Prosser, 24-VI-96 on Pyrus malus (96G279*). 

Hayhurstia atriplicis (L.) [on Chenopodium album]: ID, LataH—Moscow, 28-IX-96 (96T561); WA, ASOTIN— 
Anatone 28-VI-96 (96T091); De Spain Springs, 31-VII-96 (961358); BENTON—WSU-Prosser, 20-VIII-96, 
& 3,18,25-VII-97 (96G509, 97F190, 97F238, 97F262); KLickirat—Bickleton 18-VI-97, 7-VII-97 
(97G111, 97G118, 97G239); nr Cleveland, Dot & Brannon Rds, 27-VI-97, 7-VII-97 (97G143, 97G231); 
WHITMAN—Pullman, 16 & 30-VII-96, 7-VIII-96 (961302, 961360, 96T361+, 96T417); YAKIMA [*]— 
Grandview, 22-VII-95, 17-V-96, 3-VI-96, 8 & 30-VII-96, 29-VI-97 (95G367, 96G067, 96G131, 96G327, 
96G430. 97G181): nr Bickleton, 29-VIII-95 (95G524); Glade Cr, 7-VII-97 (97G218); Bickleton Hwy & 
County Line Rd, 7-VI-97 (97G236); Wenas Cr & Maloy Rd, 11-VI-97 (97G065); Wenas Lk, 17-VI-97 
(97G091); YIR, Signal Peak Guard Station, 19-I[X-96 (96G578). 

Illinoia corylina (Davidson): WA, WHITMAN—Pullman, 14-VIII-96 on Aquilegia sp. (96T442). 

Lipaphis erysimi (Kaltenbach): WA, BENTON—WSU-Prosser, 30-IX-96, on Brassica napus (96G6217); 
WHITMAN—Pullman, 3-VII-96 on Capsella bursa-pastoris (9611667). 

Macrosiphum euphorbiae (Thomas): ID, LATAH—Moscow, 7-X-96, on Solanum sp. (961577); WA, PIERCE 
[on Epilobium angustifolium* }]. 

Macrosiphum nr euphorbiae: WA, YAKIMA—YIR, Mill Cr Guard Station, 6-VI-95 on Lomatium nudicaule 
(95G115). 

Macrosiphum sp.: WA, ASOTIN [on Triticum aestivum*). 

Myzus persicae (Sulzer): WA, BENTON [on Raphanus sativum*|—23-VI-96 on Chrysanthemum sp. 
(96G388+): WSU-Prosser, 29-VII-96 on Carthamus tinctorius (96G428); 31-VIII-95, 1-X-97 on 
Chenopodium album (97G476); 27-X-95 on Malva neglecta (95K212+); 1-[X-95 on Solanum tuberosum 
(95G525, 95G526): 6-VII-95 on Sonchus oleraceus (95A026+); Prosser, 23-VIH-96 on Chrysanthemum sp. 
(96G388); SKAGIT [on Brassica oleracea*], SNOHOMISH—Monroe, 24-VII-96 on Pisum sativum (9615947); 
WHITMAN—Pullman, 16-VII-95, 3-VII-95 on Raphanus sativum (951248, 95T407), 22-VII-95 on Solanum 
lycopersicon (95T299+), & 3-VIII-95 on Spinacia oleracea (95T408); Ridpath, 6 & 22-VII-95 on 
Amsinckia retrorsa (95G295+).. 

Myzus sp.: WA, WHITMAN [on Forsythia sp.*]. 

Phorodon humuli (Schrank) [on Humulus lupulus unless otherwise indicated |]: WA, BENTON [*]—WSU- 
Prosser, 9-IX-95 (95K154), & 27-X-95 on Lamium amplexicaule (95K213+); YAKIMA—Grandview, 8-VI- 
95 (95G145). 

Rhopalosiphum insertum (Walker): WA, DouGLas—Orondo, 28-V-96 on Pyrus malus (96RO003). 

Rhopalosiphum maidis (Fitch): WA, BENTON [on Panicum milliaceum* |—WSU-Prosser, 18-XI-95 & 20-I-96 
on Capsella bursa-pastoris (95G750+, 96GO03+), & 2-IX-96 on Hordeum vulgare (96G655). 

Rhopalosiphum padi (L.) [on Triticum aestivum unless otherwise indicated]: WA, BENTON [on Hordeum 
vulgare & Triticum aestivum*|—Prosser, 20-VIII-96 on Zea mays (96G510); WSU-Prosser, 1-X-95, 24- 
VI-96 (95G668, 96G272): Cemetery Rd & Hwy 241, 12-X-95 (95G690); DouGLAs—Waterville, 12-VIII- 
95 (95B016, 95B018); YAKIMA [*]—Grandview, 22-X-97 (97G490). 

Sitobion avenae (FE) [on Triticum aestivum unless otherwise indicated]: WA, BENTON [*]—Sellards & Travis 
Rds, 25-VI-95 (95G253); Prosser, 30-VI-95, 17-VII-95, 24-VI-96, 1 & 8-VII-96, (95G285, 95G352, 
95G354, 95G356, 95G358, 96G271, 96G312, 96G330); WSU-Prosser, 22,30-VI-97, 3,11,18,25-VII-97 
(assorted collections between 97F183 & 97F277), & 18-VII-97 on Setaria lutescens (97G243); nr 
Whitstran, 17-VII-95, 8-VII-96 (95G355, 96G328); FRANKLIN—Star School, 6-VII-95 (95G289); Kahlotus, 


VOLUME 102, NUMBER 3 705 


6-VII-95 (95G290); KLickiTaT [*]—Bickleton, 9-VII-95 (95G318); WHITMAN [*]—Rosalia, 16-VII-96 
(96T294); YAKIMA—Grandview, 5-VII-95 (95G286). 

Uroleucon ivae (Robinson): WA, YAKIMA—Grandview, 9-VII-95, on Iva xanthifolia (95G313). 

Uroleucon sp. YAKIMA—Grandview, 18-I-96, on Brassica oleracea (96GO001). 


Genus EPHEDRUS Haliday 


Ephedrus californicus Baker 

Acyrthosiphon lactucae (Passerini) [on Lactuca serriola]: WA, Asotin [*]; CHELAN—Chelan SP, 11-[X-97 
(97K038); YAKIMA—YIR, Signal Peak Guard Station, 19-[X-95 (95G621). 

Aphis fabae Scopoli: WA, AsoTiIn—Field Springs SP, 18-VII-95 on Cirsium arvense (951272). 

Aphis helianthi Monell: MT, SANDERS—nr Thompson Pass, 29-VII-95 on Ligusticum verticillatum (95G460); 
WA, CHELAN—Stevens Pass, 16-IX-95 on Epilobium angustifolium (95K1347); GARFIELD—Stentz Springs, 
22-VIII-95 (95T452); Kitriras—Lost Lk, 24-VII-95 on Epilobium angustifolium (95G399*7), & 11-IX-95 
on Heracleum lanatum (95G572); 2-[X-95 on Lomatium sp. (95G5357+); LEwis—RNP, Bench Lk, 8-[X-95 
on Ligusticum sp. (95K113); PIlERCE—RNP, Tipsoo Lk, 11-[X-96 on Osmorhiza sp. (96G535+); YAKIMA— 
RNP, Dewey Lk Trail, 11-[X-96 on Ligusticum sp. (96G5527+); YIR, 25-V-96 on Lomatium dissectum 
(96K052), Mt Adams Lk, 25-VII-95 on Osmorhiza occidentalis (95G428), & Windy Point, 19-[X-95 on 
Xerophyllum tenax (95G635). 

Aphis holodisci Robinson [on Holodiscus discolor unless otherwise indicated]: WA, SPOKANE—Mt Spokane 
SP, 27-VII-95 (951338); Manito Pk, 11-VII-96 (96T260); YAKIMA—Clear Lk, 15-VI-96 (96K082); Bird 
Cr Meadows Rd & FR822, 26-VI-95 on Physocarpus malvaceus (95A020, 95A021). 

Aphis nr oenotherae Oestlund: WA, GARFIELD—Pataha Cr Rd, 7-I[X-95 on Epilobium sp. (951541). 

Aphis varians Patch [on Epilobium angustifolium]: ID, SHOSsHONE—Thompson Pass, 29-VII-95 (95G4687); 
WA, CHELAN—Stevens Pass, 16-IX-95 (95K1347); Kitriras—Lost Lk, 24-VII-95 (95G3997); PIERCE— 
Naches Pass, 25-IX-97 (97G460); RNP, Bench Lk, 8-IX-95 (95K114). 

Aphis nr varians Patch: WA, Kittrras—Lost Lk, 24-VII-95 on Epilobium angustifolium (95G399*). 

Aphthargelia symphoricarpi (Thomas) [on Symphoricarpos sp.\: WA, Kittitas [*]; SPOKANE—nr Peone, 
Cooper Rd & Hwy 206, 29-VI-95 (95T178); YAkIMA—nr Rimrock Lk, 9-VI-95 (95A004). 

Brachycaudus helichrysi (Kaltenbach): WA, ASoTIN—nr FR 40 & FR44, 1-VIII-95 on Achillea millefolium 
(O5T3 827): 

Braggia eriogoni (Cowen): WA, KittirAs—Taneum Lk, 11-IX-97 on Epilobium umbellatum (97G428). 

Brevicoryne brassicae (L.): WA, FRANKLIN—Eureka, 14-V-96 on Brassica napus (96G040). 

Capitophorus elaeagni (del Guercio): WA, YAKIMA—FR 1010, 8 mi S of Tieton River, 14-IX-95 on Cirsium 
vulgare (95G596). 

Cavariella aegopodii (Scopoli) [on Ligusticum grayi]: WA, PIERCE—RNP, Tipsoo Lk, 5-IX-97 (97G3637); 
YAKIMA, YIR, Windy Point, 19-IX-95 (95G6427+). 

Cavariella pastinacae (L.): WA, AsoTin—Asotin Cr Rd, 11-VII-95 on Oenothera sp. (951196). 

Chaetosiphon fragaefolii (Cockerell) [Rosa sp.]: ID, BOUNDARY [{*]; WA, PEND OrREILLE—Le Clerc WLA, 
3-VIII-95 (95T398). 

Diuraphis noxia (Kurdjumov): WA, Asotin—Anatone, 8-VIII-96 on Hordeum vulgare (961598); Asotin Cr 
WLA, 3-X-96 on Triticum aestivum (96T565). 

Ericaphis wakibae (Hottes): WA, YAKIMA—Clear Lk, 15-VI-96 on native Rosa sp. (96KO0857%). 

Hyperomyzus lactucae (L.): WA, KLickiIrAt—Rock Cr & Columbia River, 28-IV-95 on Sonchus oleraceus 
(95GO0117). , 

Hyperomyzus nigricornis (Knowlton) [on Hieracium sp. unless otherwise indicated]: ID, LATAH—Moscow 
Mtn 13-VII-96 on unknown plant (96T264); 3,15-VIII-96, on Compositae (961399, 96T446); MT, 
SANDERS—nr Thompson Pass, 29-VII-95 (95G462+); WA, GarFiELD—Alder Thicket, 21-IX-95 (95T587); 
KittirAs—Manastash Rd, 25-VII-96, 16-VII-96 (96G415, 96G490); Manastash Cr & FR125, 16-VIII-96 
(96G493); Ellensburg, 23-VI-96 on Ribes sp. (96G267); Lost Lk, 6-VIII-97 (97G344); Klickitat— 
Cleveland, 16-VI-96 on Madia minima (96K109); 7-VII-97 (97G229); PIERCE—RNP, Tipsoo Lk, 11-IX- 
96 (96G541), & 5-IX-97 on Agoseris aurantica (97G3847+); SPOKANE—Mt Spokane SP, 24-VIII-95 on 
Compositae (95T486); Mt Spokane Hwy, mi marker 18.5, 22-VIII-96, on Compositae (96T487, 96T488); 
WHITMAN—Kamiak Butte, 7-VII-96, on unknown plant (96T205); Pullman, 10-VIII-95 on Lapsana 
communis (95T436+); YAKIMA—Ravens Roost, 28-VII-97 on Hieracium albertinum (97G296); Wenas Lk, 
30-V-96 on Ribes aureum (96G123); YIR, Signal Peak, 19-I[X-95 (95G622) & Fish Lk (95G637) on 
Agoseris aurantica, Delaney Springs, 25-VII-95 on Hieracium cynoglossoides (95G406), & Goat Butt, 19- 
IX-96 (96G585); 

Hyperomyzus sp.: WA, Kittrras—S. Fork Manastash Cr, 24-VII-95 on Hieracium albiflorum (95G388). 


706 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Illinoia sp.: WA, GARFIELD—FR 160 & FR 40, 21-[X-95 on Vaccinium sp. (95T596). 

Macrosiphoniella ludovicianae (Oestlund) [on Artemisia ludoviciana|: WA, YAKIMA [*]; YIR, Randell 
Springs, 16-V-96 (96G049¥*). 

Macrosiphoniella tanacetaria (Kaltenbach): WA, CHELAN—Stevens Pass, 16-IX-95 on Tanacetum vulgare 
(95K133). 

Macrosiphum albifrons Essig [on Lupinus sp.): WA, KittirAs—Buck Meadows, 9-VII-96 (96G338); 
KLickITAt—Trout Lk, 11-VII-95 (95K068); LEwis—RNP, Stevens Canyon, 23-VII-96 (96G364); PIERCE— 
RNP, Tipsoo Lk, 11-IX-96 (96G547) & 5-IX-97 (97G362); SPOKANE—Manito Pk, 11-VI-96 (96T033); nr 
Peone, 2-VII-96 (96T098); YAKIMA—American River, 5-I[X-97 (97G376). 

Macrosiphum californicum (Clarke): WA, Kittrras—Murray WLA, S Fork Manastash River, 11-[X-95 on 
Salix sp. (95G3917). 

Macrosiphum creelii Davis: WA, Kittiras—Manastash Cr Rd, mi marker 8, 16-VIII-96 on Vicia sp. (96G492). 

Macrosiphum euphorbiae (Thomas): ID, LATAH—Moscow Mtn, 6-VII-96 on unknown plant (96T181); 6-VII- 
96 on Compositae (96T185, 961192); MT, SANDERS—nr Thompson Pass, 29-VII-95 on Hieracium sp. 
(95G462+); WA, AsoTIN—De Spain Springs, 11-VU-96 on unknown plant (96T252); nr FR 40 & FR44, 
1-VIII-95 on Achillea millefolium (95T382+); BENTON — WSU-Prosser, 27-X-95 on cultivated Rosa sp. 
(95K220+); GARFIELD—Stentz Springs, 1-VII-95 on Arnica latifolia (95T370); KING — Issaquah, 4-VI-96 
on native Rosa sp. (96G1457+); Kittiras [on Rosa sp.*]|—Buck Meadows, 9-VII-96 on native Rosa sp. 
(96G335) & 16-VIII-96 on Silene sp. (96G497); Lost Lk Trail, 6-VIII-97 on Luina nardosmia (97G327); 
Stampede Pass, 8-VIII-96 on Castilleja sp. (96K1407+); KLIckIrat—Cleveland, 7-VII-97 on Lupinus sp. 
(97G230); LEwis—RNP, Stevens Canyon, 23-VII-96 on Crepis sp. (96G357); PEND OREILLE—Cusick, 9- 
VII-96 on native Rosa sp. (961223); SKAMANIA—Twin Falls CG, 4-VIII-95 on Potentilla sp. (95K0987); 
WHITMAN—Kamiak Butte, 7-VII-96 on Compositae (96T200); Pullman, 10-VIII-95 on Lapsana communis 
(95T4367) & 18-VI-96 on cultivated Rosa sp.(96T037); YAKIMA [on Ligusticum grayi+* |—Clear Lk, 15- 
VI-96 on native Rosa sp. (96K0857); N. Fork Tieton River, 15-VI-96, on Hieracium albertinum (96KO094); 
Buckhorn Meadows, 14-IX-95 on Trautvetteria caroliniensis (95G597); YIR, Bird Cr Meadows, 26-VI- 
95, on Osmorhiza chilensis (95A014). 

Macrosiphum nr euphorbiae (Thomas): WA, YAKIMA—N. Fork Wenas Cr, 30-V-96 on Arabis sp. (96G129). 

Macrosiphum gaurae (Williams): WA, WHITMAN—Colton, 19-IX-95 on Gaura parviflora (95T585). 

Macrosiphum pallidum (Oestlund): WA, SKAMANIA—Twin Falls CG, 4-VIII-95 on Potentilla sp. (QSKO987#). 

Macrosiphum rosae (L.) [on Rosa sp. unless otherwise indicated]: OR, BENToN—Corvallis, 18-V-93 on 
unknown plant, A. Jensen collector (93J002); WA, BENTON—-WSU-Prosser, 27-X-95 (95K2207); KING— 
Issaquah, 4-VI-96 (96G145+); SPOKANE—Manito Pk, 19-IX-96 (96T540); YAKIMA [*]—Grandview, 13 & 
21-V-97 (97G021+, 97G037). 

Macrosiphum valerianae (Clarke): WA, Kirtiras—Lost Lk, 24-VII-95 on Epilobium angustifolium 
(95G3997x). 

Metopolophium dirhodum (Walker): WA, BENTON—-WSU-Prosser, 27-X-95 on Conyza canadensis 
(95K2167+#). 

Myzus cerasi (FE): WA, BENTON—Prosser, 28-V-96 on Prunus sp. (96G091); WHITMAN—Pullman, 5-VII-96 on 
P. avium (9671173). 

Myzus persicae (Sulzer): WA, YAKIMA—Cliffdell, 23-X-97 on Linaria dalmatica (97G496). 

Nasonovia ribisnigri (Mosley): WA, WHITMAN—Pullman, 5-VII-96 on unknown plant (96T177). 

Nasonovia (Kakimia) sp.: WA, WHITMAN—Pullman, 10-VIII-95 on Lapsana communis (9574367). 

Obtusicauda artemisiphila (Knowlton & Allen): WA, KLICKITAT [on Artemisia tridentata*]. 

Obtusicauda coweni (Hunter): WA, YAKIMA [on Artemisia tridentata* |. 

Obtusicauda sp.: WA, KLICKITAT [on Artemisia tridentata*}]. 

Pleotrichophorus nr amsinckii Richards: WA, GARFIELD—Alder Thicket CG, 14-IX-95 on Amsinckia sp. 
(95T562). 

Pleotrichophorus pseudoglandulosus (Palmer): WA, YAKIMA—YIR, Randell Springs, 16-V-96 on Artemisia 
ludoviciana (96G049¥+¢). 

Rhopalosiphum insertum (Walker): WA, Kittiras—Ellensburg, 17-V-96 on Crataegus columbiana (96G066). 

Rhopalosiphum maidis (Fitch): WA, BENTON—WSU-Prosser, 18-XI-95 on Capsella bursa-pastoris 
(95G750F#). 

Rhopalosiphum padi (L.): WA, BENTON—-WSU-Prosser, 18-XI-95 on Capsella bursa-pastoris (95G7507#). 

Sitobion avenae (F.): WA, Kinc—Stampede Pass on Dactylis glomerata (96G442); Kirtiras—Manastash Rd 
& FR 112, 9-VII-96 on D. glomerata (96G346); PIERCE—Graham, 25-V-96, on Gramineae (96T001); RNP, 
Tipsoo Lk, 5-IX-97 on Agoseris aurantica (97G384+);YAKIMA—Blowout Mtn, 9-IX-97 on D. glomerata 
(97G401). 


VOLUME 102, NUMBER 3 707 


Uroleucon nr arnesense Robinson: MT, SANDERS—18 mi E of Thompson Pass, 29-VII-95 on Solidago sp. 
(95G457). 

Uroleucon erigeronense (Thomas): WA, BENTON—-WSU-Prosser, 27-X-95 on Conyza canadensis (95K2167); 
CHELAN—Leavenworth, 16-IX-95 on Conyza canadensis (95K141); KitriraAs—Lost Lk, 2-IX-95, on 
Anaphalis margaritacea (95G532) & 6-VIII-97 on Aster sp. (97G346); Manastash Rd & FR 112, 11-IX- 
95 on Achillea millefolium (95G563), on Anaphalis margaritacea (95G5617) & Cirsium arvense 
(95G564+); PIERCE—RNP, Tipsoo Lk, 5-IX-95 (95K106) & 8-IX-97 (97G3647+) on Aster sp.; WHITMAN— 
Pullman, 10-IX-96 on unknown plant (96T530); YAKiImMA—Hells Crossing CG, 11-IX-96 on Aster sp. 
(96G524+); YIR, White Deer Cr, 11-VII-95 on Anaphalis margaritacea (95G326), & Signal Peak, 19-IX- 
95 on Achillea millefolium (95G6207). 

Uroleucon nr erigeronense (Thomas): WA, OKANOGAN—Bridgeport SP, 9-IX-97 on Conyza canadensis 
(97K023). 

Uroleucon ivae Robinson: WA, YAKIMA—Grandview, on Iva xanthifolia (96G431). 

Uroleucon katonkae (Hottes): WA, YAKIMA—Hells Crossing CG, 11-[X-96 on Aster sp. (96G5247). 

Uroleucon russellae (HRL) [on Anaphalis margeritacea|: MT, SANDERS [*]; WA, CHELAN—Chatter Cr, 1- 
VIII-95 (95G484); Kitriras—Manastash Rd & FR112, 11-[X-95, on Anaphalis margaritacea (95G5617*); 
SKAMANIA—June Lk, 4-VIII-95 (95K101); SPOKANE—Mt Spokane SP, 24-VIII-95 (95T494). 

Uroleucon sonchi (L.) [on Sonchus oleraceus unless otherwise indicated]: WA, CLARK—-WSU-Vancouver, 21- 
IX-95 (95K172); KLickITAT—Rock Cr & Columbia River, 28-IV-95 (95G0O117); PlERCE—Graham, 8-IX- 
96, on Lactuca sp. (961527); WHITMAN—Pullman, 9-IX-96 on Lactuca sp. (96T529); YAKIMA—Yakima 
Arboretum, 26-IX-95 (95G6587); YIR, Satus, 1-VI-96, on Sonchus sp. (96K064). 

Uroleucon spp.: ID, BENEWAH—McCroskey SP, 31-VIII-95 on Adenocaulon bicolor (95T518, 951527, 
95T536); WA, AsoTIN [on Dipsacus sylvestris*]; CHELAN—Chiwawa Canyon, 3-VIII-96 on A. bicolor 
(96R044); Kitriras—Lost Lk, 6-VIII-97 on Achillea millefolium (97G318); S. Fork Manastash Cr, 24-VII- 
95 on Hieracium sp. (95G389); Kiickirat—nr Bickleton, Box Springs, 27-VI-97 on Balsamorhiza 
sagittata (97G154); PEND OREILLE—Sullivan Lk, 3-VIII-95 (95T391); Tiger Meadows, 9-VII-96 on Senecio 
sp. (961220); PIERCE—RNP, Tipsoo Lk, 5-IX-97 on Anaphalis margaritacea (97G367); SPOKANE—Mt 
Spokane SP, 24-VIII-95 on Centaurea pratensis (95T477); WHITMAN [on Lactuca serriola*|; YAKIMA [on 
Aster sp.*|—Union Gap, 23-X-95 on Centaurea maculosa (95G720); YIR, Signal Peak, 19-IX-95 on 
Grindelia sp. (95G618). 

Wahlgreniella nervata (Gillette): WA, YAKIMA—Grandview, 13-V-97 on cultivated Rosa sp. (97G0217¢). 

Undetermined spp.: WA, Spokane—Mt Spokane SP, 24-VIII-95 on Epilobium sp. (95T490); WHttmMAN— 
Pullman, 25-VI-95 on Lapsana communis (951332); YAKIMA—YIR, Mabton Rd, 16-V-96 on Crepis 
atrabarba (96G050). 


Ephedrus chaitophori Gardenfors 
Chaitophorus populifolii (Essig) [on Populus trichocarpa]: WA, Kitt1ras—Manastash Cr Rd, 5 mi marker, 
16-VIII-96 (96G485); YAKIMA—Wenas Lk, 11-VI-97 (97G062). 


Ephedrus clavicornis Pike & Stary 

Aphis helianthi Monell: WA, PEND OREILLE [on unknown Umbelliferae+*]; WHITMAN [on Helianthus annuus*]. 

Aphis nr oenotherae: WA, GARFIELD, Kittitas, & Lewis [on E. angustifolium*]; YAKIMA [on E. spp. & E. 
minutum* |—Wenas Cr, 13 VIII-96 (96G467) on Epilobium sp. 

Aphis varians Patch [on Epilobium angustifolium]: MT, SANDERS, W. Fork Thompson River, 26-VII-94 
(94R029); WA, Lewis [*] & YAKIMA [*]. 

Aphthargelia symphoricarpi (Thomas): WA, YAKIMA [on Symphoricarpos sp.*}. 

Dysaphis plantaginea (Passerini): WA, DouGLAS [on Pyrus malus*]. 

Illinoia maxima (Mason): WA, SPOKANE [on Rubus parviflorus*]. 

Prociphilus sp.: WA, YAKIMA [on Amelanchier alnifolia*]. 

Sitobion avenae (EF): WA, LEwis [on Agropyron caninum*]. 

Undetermined spp. [on Epilobium sp. unless otherwise indicated]: CANADA, BritIsH CoLuMBIA [from 
sweepings*]; WA, GARFIELD [*], Kittitas [*], & YAKIMA [*]. 


Ephedrus incompletus (Provancher) 
Ovatus crataegarius (Walker): ID, BONNER—Sandpoint, 10-IV-97 on Origanum sp. (97T003). 


Ephedrus lacertosus (Haliday) 
Aspidophorodon nr longicauda (Richards): WA, SKAMANIA—nr Keenes Horse Camp, 11-VII-95 on Lupinus 
sp. (95K065+#). 
Ericaphis gentneri (Mason): WA, CHELAN—Stevens Pass, 16-IX-95 on Spiraea densiflora (95K138). 
Macrosiphum albifrons Essig: WA, SKAMANIA—nr Keenes Horse Camp, 11-VII-95 on Lupinus sp. (95K066). 


708 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Macrosiphum euphorbiae (Thomas): WA, YAKIMA—YIR, Old Maid Canyon, 25-VII-95 on Cynoglossum 
grande (95G440). 

Macrosiphum sp.: WA, CHELAN—Chatter Cr, 1-VII-95 on Oplopanax horridum (95G4887). 

Macrosiphini aphids [immatures]: WA, YAKIMA—Morse Cr, 24-VII-97 on Achlys triphylla (97G255). 

Metopolophium sp.: WA, SKAMANIA—nr Keenes Horse Camp, 11-VII-95 on Lupinus sp. (9SKO065**+). 

Sitobion sp.: OR, BENTON—Mary’s Peak, 14-[X-93 on Dicentra formosa, A. Jensen collector (93J006). 


Ephedrus persicae Froggatt 
Brachycaudus cardui (L.): WA, SkAGiIr—La Conner, 5-VI-96 on Cirsium vulgare (96G189). 
Brachycaudus helichrysi (Kaltenbach): WA, SKAGITr—La Conner, 5-VI-96 on Prunus salicina (96G196). 
Undetermined sp.: WA, WHITMAN—Pullman, 3-VIII-96 on Pyrus malus (96T042). 


Ephedrus spp. 

Aphis coweni Palmer: WA, KittiTAs [on Veratrum viride*]. 

Aphis varians Patch: MT, SANDERS, Thompson Pass, 27-VII-94 on Epilobium angustifolium (94R040). 

Chaetosiphon fragaefolii (Cockerell): WA, STEVENS—Little Pend Oreille National WLA, 9-VII-96 on native 
Rosa sp. (96T2097#). 

Diuraphis noxia (Kurdjumoyv): WA, BENTON—WSU-Prosser, 15-V-97 on Triticum aestivum (97F045). 

Ericaphis scammelli (Mason): WA, STEVENS—Little Pend Oreille National WLA, 9-VII-96 on native Rosa 
sp. (96T209+£). 

Hyperomyzus nigricornis (Knowlton): ID, LATAH—Moscow Mtn, 3-VIII-96 on Compositae (96T402). 

Macrosiphum nr euphorbiae (Thomas): WA, YAKIMA—N. Fork Tieton River, 27-VI-96 on Lilium 
columbianum (96G299). 

Macrosiphum rhamni (Clarke): ID, SHOSHONE—Mutrray, 17-VI-95 on Rhamnus purshiana (95G192). 

Prociphilus sp.: WA, WHITMAN—Pullman, 13-VI-96 on unknown plant (96T029). 

Undetermined sp.: WA, GARFIELD—Spruce Springs, 14-I[X-95 on Cirsium vulgare (951571). 


Genus EUAPHIDIUS Mackauer 


Euaphidius cingulatus (Ruthe) 

Pterocomma beulahense (Cockerell): WA, KinG—Stampede Pass, 8-VIII-96 on Populus trichocarpa 
(96G445+); WHITMAN—Pullman, 9-X-96 on Populus sp. (96T584). 

Pterocomma bicolor (Oestlund) [on Salix sp. unless otherwise indicated]: WA, KINGc—Stampede Pass, 8-VIII- 
96 on Populus trichocarpa (96G445+); KLICKIrAT—nr Bickleton, Pine Cr, 18-VI-97, on P. trichocarpa 
(97G113); PreRcE [*]; SKAMANIA—Council Lk, 4-VIII-95, (95K096); WHITMAN—Pullman, 12-IX-95 
(95T558+); YAKIMA—Wenas Lk, 17-VI-97 (97G093). 

Pterocomma sanguiceps Richards: WA, KirrirAs—Ellensburg, 10-IV-95 on Salix (95G0047). 

Pterocomma smithiae (Monell) [on Salix sp.]: WA, KittrrAs—Ellensburg, 10-IV-95 (95G004+); WHITMAN— 
Pullman, 12-IX-95 (95T5587). 

Pterocomma sp.: OR, MULTNOMAH [on Salix sp.*]; WA, YAKIMA—YIR, Signal Peak, 19-IX-95 on Salix sp. 
(95G619). 

Undetermined sp.: WA, AsoTIN—De Spain Springs, 26-VI-96 on Salix sp. (96T057). 


Euaphidius setiger Mackauer 
Periphyllus californiensis (Shinji): OR, BENTON—Corvallis, McDonald State Forest, 1-V-93 on unknown plant, 
A. Jensen collector (93J001). 
Periphyllus lyropictus (Kessler): WA, BENTON [on Acer platanoides *|; GRANT—Sun Lakes SP, 8-IX-97 on 
A. sp. (97K010); YAKIMA—Grandview, 13-VI-96 on A. sp. (96G243). 
Periphyllus sp.: WA, SPOKANE—Spokane, Manito Pk, 19-I[X-96 on Acer sp. (96T539). 


Genus HARKERIA Cameron 


Harkeria rufa Cameron 
Nasonovia houghtonensis (Troop): OR, BENTON—Corvallis, McDonald State Forest, 15-IV-94 on Oemleria 
cerasiformis, A. Jensen collector (94J002). 


Genus LYSAPHIDUS Smith 


Lysaphidus adelocarinus (Smith) 
Aphis canae Williams: WA, AsoTiIn—Asotin Cr Rd, 13-VI-95 on Artemisia tridentata (95T089). 


VOLUME 102, NUMBER 3 709 


Aphis (Zyxaphis) sp.: WA, KirrirAs—Frost Meadows, Manastash Rd, 11-IX-97 on Artemisia tridentata 
(97G435). 

Artemisaphis artemisicola (Williams): WA, FRANKLIN—Kahlotus Hwy, 28-V-96 on Artemisia tridentata 
(96G096). 

Epameibaphis atricornis G&P [on Artemisaphis tridentata]: WA, BENTON [+*]—ALE, 19-V-95 (95G0727); 
YAKIMA [{*]—nr Mabton, 9-V-95 (95G0317); Selah Cr & Hwy 821, 15-V-95 (95G0677). 

Flabellomicrosiphum knowltoni Smith [on Artemisaphis tridentata]: WA, BENTON [t*]; YAKIMA—YIR, Ft. 
Simcoe, 30-V-96 (95G0917). 

Flabellomicrosiphum tridentatae (Wilson): WA, KLICKITAaT—Alderdale Cr & Columbia River, 28-IV-95 on 
Artemisaphis tridentata (95GO137*¢#). 

Flabellomicrosiphum sp.: WA, KLICKITAT [on Artemisaphis tridentatay*]. 

Microsiphoniella artemisiae (Gillette): WA, AsotTiIn—Asotin Cr HMU, 5-VII-96 on Artemisia ludoviciana 
(96T141). 

Obtusicauda coweni (Hunter): WA, AsoTiIn—Asotin Cr HMU, 7-V-96 on Artemisia tridentata (96T 1427). 

Obtusicauda sp. [on Artemisia tridentata): WA, KLICKITAT [*] & YAKIMA [*]. 

Pleotrichophorus nr amsinckii Richards: WA, BENTON [*]—ALE, 8-VI-95 on Cryptantha sp. (95G1477). 

Pleotrichophorus palmerae (Knowlton): WA, KLickirat—nr Alderdale, 9-V-95 on Chrysothamnus sp. 
(95G0267). 

Pleotrichophorus sp.: WA, GARFIELD—Alder Thicket, 21-IX-96 on Amsinckia sp. (95T5927); KittiraAs [on 
Artemisia tridentata}*]; KLICKITAT—nr Alderdale, 9-V-95 on Chrysothamnus nauseosus (95G0267). 

Pseudoepameibaphis essigi Knowlton & Smith: WA, KLICKITAT [on Artemisia tridentata}*]. 

Pseudoepameibaphis glauca G&P [on Artemisia tridentata): WA, Kittiras [*+*]—Taneum Rd, 11-IX-95 
(95G5867); YAKIMA—YIR, Goat Butte, 9-X-96 (96K169). 

Pseudoepameibaphis tridentatae (Wilson) [on Artemisia tridentata]: WA, BENTON [*]—ALE, 19-V-95 
(95G072+); KittiTAs [*]; KLickiTat [+*]—Alderdale Cr & Columbia River, 28-IV-95 (95G0137); YAKIMA 
[+*]—Naught Rd, 18-VI-97 (97G120); nr Mabton, 9-V-95 (95G0317+); Selah Cr & Hwy 821, 15-V-95 
(95G067+); YIR, Ft. Simcoe, 30-V-96 (95G0917). 


Lysaphidus ramithyrus (Smith) 
Pleotrichophorus nr wasatchii (Knowlton): WA, KittirAs—Yakima Canyon, 11-IX-95 on Chrysothamnus sp. 
(95G5497). 
Pleotrichophorus elongatus (Knowlton): WA, LINCOLN—Lake Roosevelt, Goldsmith CG, 31-VII-97 on 
Chrysothamnus sp. (97G306). 


Lysaphidus rosaphidis (Smith) 
Aphis (Zyxaphis) sp.: WA, KirrirAs—Taneum Rd, 11-IX-95 on Artemisia tridentata (95G5857¢£). 
Chaetosiphon fragaefolii (Cockerell) [on cultivated Fragaria sp.]; WA, CLARK—Vancouver, 30-VIII-95 
(95K124); WSU-Vancouver, 21-I[X-95 (95K166, 95K170). 
Chaetosiphon nr thomasi HRL: WA, Clark—WSU-Vancouver, Greenhouse, 15-V-97 on Fragaria sp. 
(97X020). 
Pleotrichophorus sp.: KittrrAs—Taneum Rd, 11-IX-95 on Artemisia tridentata (95G5857+). 


Lysaphidus spp. 

Aphthargelia symphoricarpi (Thomas): WA, SPOKANE—Mt Spokane SP, 14-VI-95 on Symphoricarpos sp. 
(95T116). 

Chaetosiphon fragaefolii (Cockerell): WA, STEVENS—Little Pend Oreille National WLA, 9-VII-96 on native 
Rosa sp. (96T2097+#). 

Ericaphis gentneri (Mason): WA, CHELAN—Stevens Pass, 16-[X-95 on Spiraea densiflora (95K138). 

Ericaphis scammelli (Mason): WA, STEVENS—Little Pend Oreille National WLA, 9-VII-96 on native Rosa 
sp. (96T2097#). 

Macrosiphum euphorbiae (Thomas): WA, GARFIELD—FR40 nr Teal Springs, 1-VIH-95 on Penstemon sp. 
(9513797). 

Nasonovia (Kakimia) sp.: WA, SPOKANE—Mt Spokane, Scribner Rd, 29-VI-95 on Phlox sp.? (95T176). 


Genus LYSIPHLEBUS Foerster 


Lysiphlebus flavidus Gahan 
Cedoaphis incognita Hottes & Frison: WA, YAKIMA [on Symphoricarpos sp.*}. 


Lysiphlebus testaceipes (Cresson) 
Acyrthosiphon lactucae (Passerini) [on Lactuca serriola|: ID, LATAH—Moscow Mtn, 15-VIII-96 (961447); 


710 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


WA, ASOTIN [*]; BENTON—-WSU-Prosser, 5-VIII-96 (96G434); CHELAN—Chelan SP, 11-IX-97 (97K038); 
Kittitas [*]; WHITMAN—Pullman, 17-VIII-95 (95T446); YAKIMA—YIR, Signal Peak, 19-IX-95 (95G621). 

Acyrthosiphon macrosiphum (Wilson): WA, SPOKANE—nr Peone, 13-VII-95 on Ribes sp.? (95T227). 

Acyrthosiphon pisum (Harris): WA, BENTON—-WSU-Prosser, 26-IV-97 on Medicago sativa (97G014). 

Anoecia corni (F.): WA, YAKIMA—Ahtanum Mission, 26-IX-95 on Cornus stolonifera (95G6627¢£). 

Aphis armoraciae Cowen: MT, LINCOLN—Libby, 16-VI-95 on Centaurea maculosa (95G172); WA, ASOTIN— 
Anatone, 24-VII-96 on unknown plant (96T330); KittiTAs [on Mertensia paniculata *|—Reecer Cr., 20- 
VI-95 on Centaurea diffusa (95G2057); KLIicKkIrat—Bickleton, 7-VII-97 on Tragopogon dubius (97G225); 
WALLA WALLA—Prescott, 30-X-97 on Taraxacum officinale (97G509); YAKIMA—nr Hause CG, 15-VI-96 
on Achillea millefolium (96K1047); Wenas Lk, 11-VI-97 on A. millefolium (97G053). 

Aphis nr armoraciae Cowen: WA, YAKIMA—nr Parker, 28-[X-95 on Centaurea sp. (95G664). 

Aphis canae Williams: WA, AsoTtin—Asotin Cr Rd, 30-V-95 on Artemisia tridentata (95T045*$). 

Aphis ceanothi Clarke [on Ceanothus velutinus unless otherwise indicated]: MT SANDERS—13 mi E of 
Thompson Pass, 29-VII-95 (95G459); WA, Kittiras—Manastash Cr Rd, mi marker 7, 16-VIII-96 on C. 
sp. (96G487); Manastash Cr, 25-VI-9 (97G137); Quilomene WLA, 2-VIU-97 (97G213); Lost Lk, 6-VIII- 
97 (97G332); KLICKITAT—Trout Lk, 29-VI-97 on C. sanguineus (97G161); YAKIMA—YIR, nr. Mt. Adams 
Lk., 25-VII-95, 4-VIII-95 on C. sanguineus (95G409, 95K0787), Miller Point, 19-IX-95 (95G6277) on]), 
Howard Lk Rd 19-IX-95 (95G641). 

Aphis nr ceanothi Clarke: WA, YAKIMA—YIR, Signal Peak, 23-VI-95 on Ceanothus sp. (95G2317), & nr. 
Mt. Adams Lk., 19-IX-95 on C. velutinus (95G607). 

Aphis nr chrysothamni Wilson: WA, AsoTiIN—Asotin Cr Rd, 13-VI-95 on Chrysothamnus sp. (95T088). 

Aphis coweni Palmer [on Veratrum viride unless otherwise indicated]: WA, AsoTIN—De Spain Springs, 5- 
VII-96 (96T155); CHELAN—Chatter Cr, 1-VIII-85 on Pyrola asarifolia (95G4857); Kittrras—Wells 
Meadow, 25-VII-96 (96G413). 

Aphis craccivora Koch: WA, BENTON—-WSU-Prosser, 3-X-95 & 8-XI-95 on Asparagus officinalis (95G672, 
95G747); Crow Butte, 24-VI-96, on Astragalus sp. (96GO013); Kittrras—Indian John Hill, 11-X-96 on 
Kochia scoparia (96G639); KLickirat—Alderdale Rd & Hwy 14, 9-V-95 on Astragalus sp. (95G025); 
Hwy 14, 4 mi W of Alderdale Rd, 14-V-96 on Medicago sativa (96G030); WHITMAN—Pullman, 25-VII- 
95 on Capsella bursa-pastoris (951331); 2-VIII-96 on Melilotus alba (96T391); 3, 7,16 & 31-VII-96, 
2,7,16,19 & 29-VIII-96, 9-X-96 on Robinia pseudo-acacia (961133, 96T199, 96T301, 961364, 96T393, 
96T415, 961462, 96T469, 96T511 & 96T580); YAkIMA—Grandview, 30-IX-96 & 3-VIII-96 on Robinia 
pseudo-acacia (95G666, 96G433); nr Union Gap, 23-X-95 on Medicago sativa (95G721+); YIR, Yakima 
Chief & Pioneer Rds, 16-V-96 on Astragalus sp.(96GO059). 

Aphis nr epilobiaria Theobald: WA, OKANOGAN—Bridgeport SP, 9-IX-97 on Epilobium minutum (97K017*). 

Aphis fabae Scopoli [on Cirsium arvense unless otherwise indicated]: ID, LATAH—Moscow Mtn, 15-VIII-96 
(961448); 13-VII-96 on Holodiscus discolor (96T267); Moscow Mtn, nr start of Paradise Cr, 21-IX-96 
(96T542); WA, AsotTin—Field Springs SP, 18-VII-95 (95T272); Wickiup CG, 29-VIII-95 (95T504); 
Anatone, 28-VI-96 (96T083); GARFIELD—Pataha Cr Rd, 5.6 mi S of Columbia Center, 7-[X-95 (95T544); 
Columbia Center, on Philadelphus lewisii (95T430); GRANT—Sun Lakes SP, 8-IX-97 on C. sp. (97K008); 
KitrirAs—Buck Meadows, 11-IX-95 (95G590), & on C. vulgare (95G580); Frost Meadows, 11-IX-95 
(95G5837); Lost Lk, 11-IX-95 (95G556+, 95G564+, 95G571+) & on C. vulgare (95G577); 6-VIII-97, on 
Senecio sp. (97G335); Manastash Rd, 11-IX-95, 25-VII-96, & 16-VIII-96 (95G5567, 96G424, 96G478, 
[96G481 on C. vulgare]); Lost Lk Trailhead, 6-VII-97 (97G320); LEwis—RNP, Bench Lk, 23-VII-96 on 
native C. sp. (96G375); KLICKIrAT—nr Bickleton, Cemetery Rd 1 mi N of Hwy, 18-VI-97, 7-VII-97 
(97G107, 97G227); Klickitat River Hwy 142, 28-VI-97 on C. vulgare (97G155); OKANOGAN—Winthrop, 
10-IX-97 on Arctium minus (97K031); SPOKANE—Mt Spokane SP, 20 & 27-VII-95, 24-VIII-95, (95T293, 
95T340, 95T349, 951474): 13-VII-95, on Umbelliferae (95T241); 24-VIII-95 on C. ochrocentrum 
(951476, 95T481); WHITMAN—Pullman, 1-VIII-95, & 7, 18-VII-96 (95T387, 96T198, 96T323); YAKIMA— 
Buckhorn Meadows, 14-IX-95 on C. vulgare (95G600); American River, 5-IX-97 (97G373) & on 
Epilobium angustifolium (97G377+); Blowout Mtn, end of Little River Rd, 9-[X-97 (97G390); YIR, Piscoe 
Cr 25-VII-95, on Rumex crispus (95G437); Signal Peak, 19-IX-96 on C. vulgare (96G563). 

Aphis nr fabae Scopoli: ID, LATAH—Moscow Mtn, 15-VIII-96 on Rumex crispus (96T450); WA, PEND 
OREILLE—Sullivan Lk Rd & Harvey Cr, on Chrysanthemum leucanthemum (951394); PIERCE—RNP, 
Tipsoo Lk, 11-IX-96 on Osmorhiza sp. (96G531). 

Aphis filifoliae (G&P): WA, Asotin—Asotin Cr Rd, 11-VII-95 on Artemisia tridentata (95T202). 

Aphis nr frangulae Kaltenbach [on Nepeta cataria]: WA, ASOoTIN—Couse Cr Rd, 10-VIII-95 (95T424); nr 
FR40 & FR44, 5 & 19-VII-96 (96T161, 961242, 961243); De Spain Spring, 11-VII-96 (961242, 96T243). 

Aphis gossypii Glover: WA, AsoTiIN—Heller’s Bar, 30V—95 on unknown plant (95T039). 


VOLUME 102, NUMBER 3 Till 


Aphis hederae Kaltenbach: OR, BENTON—Corvallis, 21-V-91 on unknown plant (91J001). 

Aphis helianthi Monell: ID, KooTENAI—Fernan Lk, on Rumex sp. (95K030); SHOSHONE—Marble Cr 9-VI-96 
on Philadelphus lewisii (96T012); MT, SANDERS—Thompson Pass, 17-VI-95 on Heracleum lanatum 
(95G179+); 13 mi E of Thompson Pass, 29-VII-95 on Ligusticum verticillatum (95G460); WA, AsoTIN— 
Asotin Cr HMU, 29-VIII-95 on Helianthus annuus (957496); Bracken Point, 7-[X-95 on H. annuus 
(95T548); Anatone, 11-VII-95 on Lomatium sp. (95T217); De Spain Springs, 11-VII-96 on Lomatium sp. 
(961245); CHELAN—Chatter Cr, 1-VIII-95 on Oplopanax horridum (95G4887+); Stevens Pass, 16-[X-95 
(95K 1347); GARFIELD—Stentz Springs, 22-VII-96 on Osmorhiza chilensis (957452); Kittitas [on 
Heracleum lanatum*|—S Fork Manastash Cr, 24-VII-95 on Angelica sp. (95G392); Buck Meadows, 24- 
VII-95 on native Cirsium sp. (95G396); Manastash Rd, 24-VII-95 & 11-IX-95 on Heracleum lanatum 
(95G378, 95G5607); 24-VII-95 on Penstemon sp. (95G379); Lost Lk, 11-[X-95 on Heracleum lanatum 
(95G572), 2-[X-95 on Epilobium angustifolium (95G530+), & 6-VIII-97 on Angelica arguta (97G3337+); 
Manastash Cr, mi marker 5, 16-VIII-96 on Heracleum lanatum (96G483); Reecer Cr, 20-VI-95 on 
Ligusticum sp. (95G201) & 20-VI-95 on Lomatium nudicaule (95G209); Quartz Mtn, 25-VII-96 on 
Penstemon procerus (96G399) & Ligusticum grayi (96G3987+); Colockum Ridge, 2-VII-97 on Camassia 
quamash (97G196); Colockum Ridge, 1 & 2-VII-97 on Lomatium nudicaule (97G189) & Lomatium 
triternatum (97G1917+); Quilomene WLA, 2-VII-97, on Lomatium sp. (97G212); LEwis—RNP, Stevens 
Canyon, 23-VII-96 on native Cirsium sp. (96G355) & 8-IX-95 on Ligusticum sp. (95K113); OKANOGAN— 
Bridgeport SP, 9-IX-97 on Helianthus annuus (97K016); PIERCE [on Cirsium sp. & Heracleum lanatum*|— 
RNP, nr Paradise, 8-I[X-95 on native Cirsium sp. (95K122) & Xerophyllum tenax (95K121); Tipsoo Lk, 5- 
IX-97 on Ligusticum grayi (97G3637+); SKAMANIA [on Ligusticum apiifolium*]; SPOKANE—nr Peone, 13- 
VII-95 on Cornus stolonifera (95T236); Turnbull National WLA on Cornus stolonifera (95T009); Mt 
Spokane SP, 29-VI-95 on Cornus stolonifera (95T186); nr Mt Spokane, 29-VI-95 on Heracleum lanatum 
(95T171); nr Peone, 13-VII-95 on Heracleum lanatum (95T229); WHITMAN—nr Colton, Steptoe Canyon 
Rd, 19-IX-95 on Helianthus annuus (957586); Pullman, I-VIII-95 on Coriandrum sativum (9513867); 
YAKIMA [on Cirsium sp.*|—Mabton & Alderdale Rds, 23-V-96 on Lomatium sp. (96G071); Wenas Lk, 
30-V-96 & 11-VI-97 on Lomatium sp. (96G118, 97G063+) & 17-VI-97 on unknown Umbelliferae 
(97G088); American River, 5-[X-97 on Cirsium sp. (97G379), 5-[X-97 on Heracleum lanatum (97G382); 
Blowout Mtn, end of Little Naches River Rd, 9-IX-97 on Cirsium sp. (97G399) & on Xerophyllum tenax 
(97G395, 97G398); Sawmill Flats CG, 28-VII-97 on Cornus stolonifera (97G286); YIR, Lower Mill Cr, 
25-V-96 on Lomatium dissectum (96K052) & Lomatium tritematum (96K053); Piscoe Cr, 25-VII-95 on 
Ligusticum grayi (95G432); Potato Hill Guard Station, 25-VII-95 on Xerophyllum tenax (95G429): 

Aphis nr helianthi Monell: ID, BENEWAH—McCroskey SP, 31-VIII-95 on Lomatium sp. (95T517++); WA, 
KitTirAs—Manastash Cr Rd, mi marker 4, 16-VIII-96 on Cornus stolonifera (96G474); YAKIMA—Chinook 
Pass, 27-I[X-96 on Ligusticum sp. (96G604). 

Aphis hermistonii Wilson: WA, GRANT—Sun Lakes SP, 8-IX-97 on Chrysothamnus nauseosus (97KO009). 

Aphis holodisci Robinson [on Holodiscus discolor unless otherwise indicated]: MT, SANDERS—Bull River 
Valley, 16-VI-95 on Physocarpus malvaceus (95G176); WA, AsoTiIN—De Spain Springs, 26-VI-96, 5-VII- 
96 (96T059, 96T 150); Kittrras—Manastash Cr, mi marker 12, 25-VI-97 (97G133); KLickiIrAt—Trout Lk, 
11-VH-95 (95G340+); SPOKANE—Mt Spokane SP, 27-VII-95 (951338); Spokane, Manito Pk, 11-VII-96 
(961260); YAKIMA [*]—Clear Lk, 15-VI-96 (96K079, 96K082); Bird Cr Meadows Rd, FR 82 & FR 822, 
26-V95 on Physocarpus malvaceus (95A020, 95A021); YIR, White Deer Cr, 11-VII-95 (95G324) & Signal 
Peak (95G333), & nr Mill Cr Guard Station, 25-V-96 (96K047). 

Aphis lugentis Williams: WA, YAKIMA [on Senecio triangularis & Arnica sp.*]. 

Aphis lupini G&P [on Lupinus sp.]: WA, GARFIELD—Pataha Cr, 7-IX-95 (95T546); STEVENS—Little Pend 
Oreille Nat. WLA, 9-VII-96 (96T212). 

Aphis neogillettei Palmer [on Cornus stolonifera]: WA, KLICKITAT [*]; YAKIMA—Sawmill Flats CG, 28-VII- 
97 (97G285). 

Aphis nerii Boyer de Fonscolombe [on Asclepias speciosa]: WA, BENTON [*]; CHELAN—-WSU-Wenatchee, 
25-VIII-95 (95RO025); YAKIMA [*]—Sunnyside, 24-VIII-95, 1-X-95, 23-X-95, 13-VIII-96, 29-IX-96, 4, 22- 
X-97 (95G517, 95G670, 95G704, 96G452, 96G618, 97G486, 97G489); Union Gap, 23-X-95 (95G725); 
Mabton, 30-VIII-96 (96G513). 

Aphis nigratibialis Robinson [on Cornus stolonifera |}: WA, Kitt1rAs—Manastash Cr, mi marker 12, 6-VIII- 
97 (97G312); KLickiIrat—Trout Lk, 28-VI-97 (97G156); SPOKANE—Mt Spokane SP, 13-VII-95 (951239); 
YAKIMA—Chinook Pass, 28-VII-97 (97G280). 

Aphis oenotherae Oestlund: WA, OKANOGAN—Bridgeport SP, 9-IX-97 on Epilobium minutum (97K017*); 
PIERCE—RNP, nr Chinook & Cayuse Passes, 16-VIII-95 on Ribes viscosissimum (95A039). 

Aphis nr oenotherae Oestlund [on Epilobium sp. unless otherwise indicated]: WA, BENTON—Kennewick, 28- 


712 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


[X-96 (96G597); GARFIELD—Pataha Cr Rd, 7-IX-95 (95T541, 95T545); Spruce Springs, 22-VIII-95 on 
Physostegia parviflora (95T462+); OKANOGAN—Bridgeport SP, 9-I[X-97 on Oenothera strigosa (97K020); 
YAKIMA—American River, 5-IX-97 on Epilobium minutum (97G375*); Fifes Peak Viewpoint, 27-[X-96 
(96G603); Nile, 5-[X-97 (97G358); nr Rimrock, 14-[X-95 (95G594). 

Aphis pomi de Geer: WA, DouGLAs—Bridgeport, 17-VI-96 on Pyrus malus (96X007*¥£). 

Aphis rumicis L. [on Rumex crispus]: WA, BENTON [*]; YAKIMA—YIR, Satus, 1-VI-95 (96K065). 

Aphis salicariae Koch [on Epilobium angustifolium]: ID, KOOTENAI—Rathdrum, 29-VII-95 (95G443+); WA, 
AsoTIN—Anatone, 24-VII-96 (961336); CHELAN—1-VIII-95 (95G481+); Kirriras—Lost Lk, 2-IX-95 
(95G5307); OKANOGAN—Washington Pass, 10-I[X-97 (97K033); SPOKANE—nr Cheney, 23-V-95 on Cornus 
stolonifera (95K007); YAKIMA—YIR, nr. Mt. Adams Lk., 19-I[X-95 (95G6107). 

Aphis nr salicariae Koch: WA, CHELAN—Swauk Pass, 1-VIII-95 on Epilobium angustifolium (95G492*). 

Aphis sambuci L: WA, PIERCE, [on Sambucus racemosa*}. 

Aphis spiraecola Patch: WA, BENTON [on Lactuca serriola*]; CHELAN—-WSU-Wenatchee, 7-VII-95 on Pyrus 
malus (95RO18+); KitrirAs—Lost Lk, 6-VIII-97 on Angelica arguta (97G333+); YAKIMA—Sunnyside, 3- 
VI-95, on cultivated Spiraea sp. (95G109); nr Sunnyside, 28-VI-95 (95G2597). 

Aphis nr triglochinis Theobald: WA, SPOKANE—Mt Spokane SP, 29-VI-95 on native Ribes sp. (95T189). 

Aphis nr valerianae Cowen: WA, PIERCE—RNP, Mowich Lk, 25-IX-97 on Valeriana sitchensis (97G446). 

Aphis varians Patch [on Epilobium angustifolium unless otherwise indicated]: MT, SANDERS [*]: ID, 
KooTenaI—Rathdrum, 29-VII-95 (95G443+); WA, CHELAN—Stevens Pass, 16-IX-95 (95K1347); PIERCE 
[*]—RNP, nr Paradise, 8-IX-95 (95K114, 95K117), Tipsoo Lk, 5-[X-97 (97G368), & Mowich Lk, 25-IX- 
97 (97G442): Naches Pass, 25-IX-97 (97G460); SPOKANE—Mt Spokane SP, 24-VIII-95 (95T483); YAKIMA 
[*]—Green Lk, 31-VIII-94 on E. latifolium (94R140); Blowout Mtn, end of Little Naches River Rd, 9- 
IX-97 (97G404); YIR, Hussen Meadows, 19-IX-95 (95G643) & Howard Lk,19-IX-95 on Lupinus sp. 
(95G639); 

Aphis nr varians Patch [on Epilobium angustifolium]: WA, CHELAN—Dryden, 1-VIII-95 (95G4817+); LEwis— 
RNP, Bench Lk, 23-VII-96 (96G376); Stevens Cr, 23-VII-96 (96G380). 

Aphis (Zyxaphis) sp.: WA, AsoTIN—Asotin Cr Rd, 22-VIII-95 on Chrysothamnus sp. (95T469). 

Aphis sp. n.. WA, YAKIMA—YIR, Klickitat River nr Miller Point, 23-VI-95 & 11-VII-95 on Ceanothus sp. 
(95K061, 95G230) & 19-IX-95 & 19-IX-96 on C. velutinus (95G627+, 96G580). 

Aphis sp.: ID, BENEWAH—McCroskey SP, 31-VIII-95 on Cirsium vulgare (95T531); LataH—Moscow, 15- 
IX-93 on Solanum lycopersicon (93T020); WA, AsoTiIn—Anatone, 28-VI-96, on Lamium sp. (96T086); 
Schumaker Rd mi marker 1, 10-VIII-95 on Cirsium arvense (957428); 28-VI-96, on Holodiscus discolor 
(96T085); De Spain Spring, 5-VII-96, on Nepeta cataria (96T 161); 11,24-VH-96 on unknown Umbelliferae 
(961254, 961341); CHELAN—Chelan SP, 11-IX-97, on Salix sp. (97K0427); GARFIELD—Alder Thicket (FR 
40), 22-VII-95 on native Cirsium sp. (951448); Kittiras [on Pedicularis bracteosat & Senecio 
triangularis*|—Lost Lk, 6-VIII-97 on Symphoricarpos sp. (97G341+); Taneum Lk, 11-IX-97 on 
Pedicularis racemosa (97G411+); [KLICKITAT, on Rheum rhabarbarum*|—Trout Lk, 29-VI-97 on 
Amsinckia sp. (97G172); PEND OREILLE—Cusick, 9-VII-96 on Ceanothus sanguineus (961225); PIERCE— 
RNP, Tipsoo Lk, 5-IX-97 on Valeriana sitchensis (97G360); SPOKANE—Mt Spokane SP, 14-VI-95 on 
Philadelphus lewisii (957117); 24-VIII-95 on Salix sp. (95T484), 29-VI-95 on unknown Umbelliferae 
(95T183); Hwy 206 mi marker 16.3, 8-VI-95 on unknown Umbelliferae (95T070); WHITMAN—Pullman, 
1-VII-95 on Coriandrum sativum (95T386); YAKIMA [on Umbelliferae*]—nr Chinook Pass, 11-IX-96 on 
Cryptantha sp. (96G519) & on Epilobium angustifolium (96G555); Little Naches River Rd, 28-VII-97 on 
Cryptantha thompsonii (97G291); Wenas Cr & Maloy Rd, 11-VI-97 on Heracleum lanatum (97G070); 
YIR, Delaney Springs, 25-VII-95 on Holodiscus discolor (95G404), Windy Point, 19-I[X-95 on Cimicifuga 
laciniata (95G631), & Signal Peak Guard Station, 19-[X-95 on Cornus stolonifera (95G617). 

Aphthargelia symphoricarpi (Thomas) [on Symphoricarpos sp. unless otherwise indicated]: WA, KITTITAS 
[*]—Colockum Rd, mi marker 4, 1-VII-97 (97G183); Lost Lk, 6-VHI-97 (97G334); KiickiTaT [on S. 
albus*]; SPOKANE—Mt Spokane SP, 14-VI-95, 13-VII-95 (95T116, 95T242); Turnbull Nat. WLA, 16-VII- 
96 on unknown plant (961287); Fish Lk, 14-VI-97 on S. occidentalis (97K004); YAKIMA—Wenas Cr & 
Maloy Rd, 11-VI-97 (97G066); YIR, Signal Peak, 11-VII-95 (95K057). 

Boernerina variabilis Richards: WA, KirrirAs—Buck Meadows, 9-VII-96 on Alnus sp. (96G336). 

Brachycaudus cardui (L.) [on Cirsium vulgare unless indicated otherwise]: ID, LAtAH—Moscow Mtn, 3 & 
15-VIII-96 (96T401, 961449); Nez Perce—Hwy 95 & Hwy 195, 22-VII-97 on C. arvense (97J004); WA, 
AsotTin—Heller’s Bar, 11-VII-95 on C. arvense (95T212); Anatone, 24-VII-96 (961333) & on Onopordum 
acanthium (961329); Asotin, 28-VI-96 on O. acanthium (961066); Weisenfels Ridge Rd nr Snake River 
Rd) 28-VI-96 on O. acanthium (96T070); BENTON [*]—Prosser, Rotha Rd, 24-VIII-95 (95G512); 
GaRFIELD—Spruce Springs, 22-VIII-95 on C. arvense (95T457); Pataha Cr Rd, 7-IX-95 (95T542); 


VOLUME 102, NUMBER 3 TAS 


KLIckITAT—Bickleton, 18-VI-96 & 7-VII-97 on C. sp. (96K112) & C. vulgare (97G228); Brooks Memorial 
SP, 21-IX-95 on C. sp. (95K162+): SPOKANE—Mt Spokane SP 24-VIII-95 (951475, 95T482. 95T488): 
Whitman—Pullman, 1,13-VIII-96, 3-[X-96 (96T369, 96T383, 961439, 961514); YAKIMA—nr Naches Pass, 
FR 789 & FR 1914, 9-IX-97 (97G410): YIR, nr. Mt. Adams Lk., 25-VII-95 (95G405), Signal Peak, 19- 
IX-95 (95G624), Satus, 1-VI-96 on native C. sp. (96K066), & Snow Cr, 19-IX-95 (95G608). 

Brachycaudus helichrysi (Kaltenbach): WA, FRANKLIN—Ice Harbor Dam, 28-V-97 on Amsinckia sp. (96G097): 
Kittitas [on Rudbeckia_hirta*); WHITMAN—Pullman, 19-VII-96 on Tragopogon dubius (961320): 
YAKIMA—YIR, nr. Mt. Adams Lk., 4-VIII-95 on native Cirsium sp. (95KO077). 

Brachycaudus tragopogonis (Kaltenbach) [on Tragopogon dubius]: ID, SHOSHONE—Kingston, 29-VII-95 
(95G479); OR, UMaTILLA—Milton-Freewater, 11-VI-96 (96G239); WA, BENTON—-WSU-Prosser, 27-VI- 
96, 15-VII-96, 30-V-96, 14-V-97 (96A003, 96A014, 96G100, 97G028); Rotha Rd, 18-VI-96 (96G256); 
Badger Springs, 28-IX-96 (96G595): West Richland, 27-V-96 (96K056):; CHELAN—Chelan SP, 11-IX-97 
(97K040); Kittiras—Ellensburg Pass, Wenas-Ellensburg Rd, 9-VI-96 (96G209); Indian John Hill, 11-X- 
96 (96G638); KLicKITAT [*]—Bickleton, 17,18-VI-96, 27-VIII-96, 8-X-96 (96G254, 96K113, 96G512, 
96G628); Box Springs, 27-VI-97 (97G147): WHITMAN—Pullman, 1,2,7,12-VII-96, 2,9-VIII-96, 3-X-96 
(96T126, 96T132, 961197, 96T325, 961387, 961427, 961574); Yakima—Grandview, 24-V-96, 2 & 26- 
VI-96, 13-X-96, 14-V-97 (96G080, 96G130, 96G273, 96G283, 96G646, 97G022): Halfway Flat CG, 27- 
IX-96 (96G616); Hause CG, 15-VI-96 (96K 106); nr Bickleton, 18-VI-97 & 7-VII-97 (97G102, 97G222): 
Sunnyside, 1-VI-96 (96K058): Wenas Lk, 30-V-96 (96G124); YIR, Signal Peak, 19-IX-96 (96G560) & 
Goat Butte, 19-IX-96, 9-X-96 (96G583, 96G631). 

Brachycorynella asparagi (Mordvilko) [on Asparagus officinalis]; WA, BENTON [*]—WSU-Prosser, 6,25-IX- 
95 (95K 144, 95G649). 

Braggia eriogoni (Cowen): MT. GALLATIN—Big Sky, Gallatin Rd, 26-VI-96 on Eriogonum sp. (96K119): 
WA, AsoTiIN—De Spain Springs, 11,19-VII-96 on E. sp. (961246, 961317); Round Prairie Springs, 18- 
VII-96 on E. sp. (96T311): KING—Stampede Pass, 8-VIII-96 on E. compositum (96G446); Kirtiras— 
Colockum Ridge Rd, 2-VII-97 on E. compositum (97G190, 97G193); Taneum Lk Trail, 11-IX-97 on E. 
umbellatum (97G428):; KLICKITAT [on E. heracleoides*|—Bickleton (Pine Cr Rd), 3-V-95 on E. heracleoides 
(95G099); Cemetery Rd 1 mi from Hwy, 18-VI-97 on E. sp. (97G106): Cleveland, 18-VI-96 on E. 
heracleoides (96K111). 

Braggia nr eriogoni (Cowen) [on Eriogonum compositum unless otherwise indicated]: OR, UMATILLA—Harris 
Pk, 11-VI-96 (96G234); WA, Kittiras—Lost Lk, 6-VIII-97 (97G350); KLICKITAT—Roosevelt, 14-V-96 
(96G035); Bickleton, 18-VI-97 (97G110); 18-VI-97, on E. sp. (97G109): YAKIMA—Little Naches River 
Rd, 28-VII-97, on E. strictum (97G290). 

Braggia sp.: WA, KLickiIrat—nr Bickleton, Pine Cr Rd, 31-V-95 on Eriogonum umbellatum (95G100); Box 
Springs, 29-VI-97 on E. compositum (97G176): YAKIMA [on E. compositum & E. elatum (aphid det. by 
G. Remaudiére)*]—Hause CG, 15-VI-96 on E. elatum (96K105); Wenas Lk, 11-VI-97 on E. sp. (97G060); 
YIR, Klickitat River, 7-VI-95 on E. compositum (95G126). 

Braggia sp. n. [on Eriogonum elatum unless otherwise indicated]: WA, DouGLas—Orondo, 28-V-96 
(96RO002); Kirriras—Lost Lk, 2-IX-95 on E. compositum (95G529); Reecer Cr, 20-VI-95 (95G213); 
YAKIMA—Bear Cr nr Rimrock Lk, 9-VI-95 (95A006): Naught Rd, 18-VI-97 (97G104) & 7-VII-97 on E. 
compositum (97G220); Naches, Hwy 410 & Hwy 12, 28-VII-97 (97G276); Naches River, Horseshoe Bend, 
5-IX-97 (97G357); nr Rimrock Lk, 14-IX-95 on E. sp. (95G595); Wenas Lk, 30-V-96,11-VI-97 (96G114, 
97G061); Windy Point CG, 3-VI-95, 1-VII-95, 26-VI-96 (95G105, 95K040, 96G305), & 16-VIII-95 on E. 
compositum (95A031); YIR, Fort Simcoe, 6-VI-95 (95G117), Mill Cr, 19-IX-95 (95G605), Signal Peak, 
19-IX-95 (95G626), & 20 mi S of Satus Pass, 7-VI-95 on E. sp. (95G142). 

Brevicoryne brassicae (L.): WA, BENTON—WSU-Prosser, 6-VII-95 on Brassica napus (95A024). 

Capitophorus elaeagni (del Guercio): WA, YAKIMA—Buckhorn Meadows, 14-IX-95 on Cirsium vulgare 
(95G596); nr Naches Pass, FR 789 & FR 1914, 9-IX-97 on Cirsium sp. (97G408). 

Cavariella aegopodii (Scopoli): WA, Kittiras—nr Lost Lk, 6-VIII-97 on Osmorhiza chilensis (97G331) & 
2-IX-95 on Lomatium sp. (95G535+); YAKIMA—nr Wenas Lk, 17-VI-97 on Lomatium sp. (97G095+). 
Cavariella pastinacae (L.): MT. SANDERS—Thompson Pass, 17-VI-95 on Heracleum lanatum (95G179+#); 
WA, AsoTiIn—Asotin Cr Rd, 11-VII-95 on Oenothera sp. (95T196): YAKIMA—YIR, Klickitat Canyon, 23- 

VI-95 on Heracleum lanatum (95G235). 

Ceruraphis viburnicola (Gillette): WA, YAKIMA—Yakima Arboretum, 26-IV-95 on Viburnum sp. (9SGO08). 

Chaetosiphon fragaefolii (Cockerell); WA, PEND OREILLE—Le Clerc WLA, 3-VIII-95 on native Rosa sp. 
(95T398). 

Chaitophorus populicola Thomas: WA, Asotin—Asotin Cr Rd, 23-V-95 on Populus trichocarpa (95T013). 


714 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Chaitophorus viminalis Monell: WA, YAKIMA—Naches, 27-[X-96 on Salix sp. (96G601). 

Cinara chinookiana Hottes: WA, YAKIMA—Darland Mtn, 31-VIII-94 on Abies lasiocarpa (94R138). 

Cinara sp.: WA, ASOTIN—FR44 8mi S of WLA, 18-VII-96 on Pinus ponderosa (961308). 

Diuraphis noxia (Kurdjumov) [on Triticum aestivum]: WA, AsotiIN—Anatone, 10-VIII-95 (95T427); Pataha 
Cr Rd, 5.6 mi S of Columbia Center, 10-VIII-95 (951435); BENTON [*]|—WSU-Prosser, 1-IV-97, 24-VII- 
97 (97G005, 97G274); KLickirat—nr Cleveland, 7-VII-97 (97G232, 97G234); Bickleton, 29-VII-97 
(97G300); SPOKANE—Mt Spokane SP, 22-VII-95 (95T495); YAKIMA [*]—Ridge Rd, 17-VI-96 (96G250, 
96G251); Sunnyside, 24-VII-97 (97G273). 

Dysaphis plantaginea (Passerini) [on Pyrus malus unless otherwise indicated]: WA, BENTON—Prosser, 22-VI- 
97 (97G123); CHELAN—-WSU-Wenatchee, 7-VII-95 & 30-V-96 (95RO187+, 96ROO09, 96RO11+); WHITMAN— 
Pullman, 30-VI-96, on unknown plant (96T119); YAKIMA—Grandview 26,28-VI-96 (96G282, 96G308); 
nr Sunnyside, 28-VI-95 (95G2597). 

Ericaphis nr gentneri (Mason): WA, SPOKANE—nr Peone, 13-VII-95 on Crataegus sp. (95T223). 

Eriosoma americanum (Riley): WA, CHELAN—Swakane WLA, 3-VI-96 on Ulmus americanus (96R018%). 

Eriosoma lanigerum (Hausmann): WA, CHELAN—-WSU-Wenatchee, 7-VI-95 on Pyrus malus (95RO19¢%). 

Hyadaphis foeniculi (Passerini): ID, BENEWwAH—McCroskey SP, 31-VIHI-95 on Lomatium sp. (95T5177%). 

Hyperomyzus nigricornis (Knowlton): WA, YAKIMA—Wenas Lk, 30-V-96, on Ribes aureum (96G123); YIR, 
19-IX-95 on Agoseris sp. (95G622). 

Illinoia spiraeae (MacGillivray): WA, GARFIELD—Teal Springs, 1-VIII-95 on Holodiscus discolor (951376). 

Macrosiphum euphorbiae (Thomas): ID, KooTeNAI—Rathdrum, 29-VH-95 on Epilobium angustifolium 
(95G4437+); LataH—Moscow, 29-VIII-96 on Solanum sp. (96T512+); WA, AsoTIN—De Spain Springs 5- 
VII-96 on Compositae (96T159); BENTON—Prosser, 24-X-96 on Medicago sativa (96G651); KLICKITAT— 
Trout Lk, 11-VII-95 on Holodiscus discolor (95G340*); PIERCE [on Epilobium angustifolium*]. 

Macrosiphum gaurae (Williams): WA, AsoTiIN—nr Weissenfels Ridge & Snake River Rds, 28-VI-96 on 
Oenothera sp. (96T073). 

Macrosiphum nr pallidum (Oestlund): WA, YAKIMA—American River, 5-IX-97 on Epilobium minutum 
(97G375+#). 

Macrosiphum sp. [nr fuscicornis MacDougall & pallidum (Oestlund)]: WA, CHELAN—Swauk Pass, 1-VIII-95 
on Epilobium angustifolium (95G4927). 

Myzus ascalonicus Doncaster: WA, YAKIMA [on moss*]. 

Myzus cerasi (F) [on Prunus sp. unless otherwise indicated]: WA, BENTON—-WSU-Prosser, 28-V-96 (96G091); 
CHELAN—Wood Reservoir, 6-VI-96 (96R019); Chelan SP, 11-I[X-97 (97K041); WHITmMAN—Pullman, 5-VII- 
96, on P. avium (96T173). 

Myzus nr cerasi (FE): WA, WHITMAN—Pullman, 6-VII-96 on unknown plant (961411). 

Myzus persicae (Sulzer): ID, LatrAH—Moscow, 29-VIII-96 on Solanum sp. (9615127). 

Nasonovia (Kakimia) sp.: WA, SPOKANE—nr Peone, 13-VII-95 on native Ribes sp. (951235). 

Nearctaphis bakeri (Cowen): WA, BENTON—WSU-Prosser, 5-V-95 on Crataegus sp. (95G023); KLICKITAT— 
Trout Lk, 29-VI-97 on Trifolium pratense (97G170). 

Nearctaphis californica HRL: WA, KLicktrat—Brooks Memorial SP, 8VI-94 on Crataegus douglasii (94— 
100). 

Nearctaphis kachena (Hottes): WA, Kirtrras—Taneum Lk, 11-IX-97 on Pedicularis racemosa (97G4117#). 

Nearctaphis yohoensis Bradley: WA, YAKIMA—YIR, White Deer Cr, 11-VII-95 on Sorbus sp. (95KO51). 

Nearctaphis sp. 1D, LataH—Moscow, 13-VI-97 on unknown plant (97T050). 

Phorodon humuli (Schrank); WA, BENTON [on Humulus lupulus & Prunus salicina*|—WSU-Prosser, 19,25- 
IV-95, 23-V-96, 15-IV-97, on P. salicina (95G005, 95G006, 95G007, 96G075, 97G006). 

Pseudoepameibaphis tridentatae (Wilson): WA, AsoTiIN—Asotin Cr Rd, 30-V-95 on Artemisia tridentata 
(95T0457#). 

Pterocomma sp.: WA, YAKIMA—YIR, Signal Peak, 19-IX-95, on Salix sp. (95G619). 

Rhopalosiphum cerasifoliae (Fitch) [on Prunus virginiana unless otherwise indicated]: MT, LINCOLN—Libby, 
29-VII-95 (95G451); SANDERS — 20mi E of Thompson Pass, 29-VII-95 (95G454); WA, KirtrrAs—Reecer 
Cr, 20-VI-95 (95G202); KLICKITAT—E end of Colockum Pass Rd, 2-VII-97, 97G215); WHITMAN—Pullman, 
19-VI1-96, 16-VII-96 on unknown plant (96T046, 96T300); YAkIMA—Wenas Lk, Wenas Cr & Maloy Rd, 
11-VI-97 (97G069). 

Rhopalosiphum enigmae Hottes & Frison: WA, YAKIMA—Buena, 23-X-95 on Typha latifolia (95G715). 

Rhopalosiphum insertum (Walker) [on Pyrus malus unless otherwise indicated]: WA, CHELAN—Chelan, 11- 
VI-96 (96X001, 96X002); WSU-Wenatchee, 30-V-96 (96RO011+).; DouGLAs—Bridgeport, 17-VI-96 


VOLUME 102, NUMBER 3 WAS 


(96X0077+); Orondo, 25-V-95 (95RO10); 11-VI-96 (96X003, 96X004, 96X005); Bridgeport, 12,17-VI-96 
(96X006, 96X008); Kitriras—Ellensburg, 17-V-96 on Crataegus columbiana (96G066). 

Rhopalosiphum maidis (Fitch): WA, BENTON [on Echinochloa crus-galli+, P. milliaceum, & Triticum 
aestivum*|—WSU-Prosser 23-XI-94 on Panicum milliaceum (94G113), 2-XI-96 on Hordeum vulgare 
(96G655), 18-XI-95 & 20-I-96 on Capsella bursa-pastoris (96GO003+, 95G7507*). 

Rhopalosiphum nymphaeae (L.): WA, YAKIMA—Outlook, 23-X-95 on Typha latifolia (95G709). 

Rhopalosiphum padi (L.) [on Triticum aestivum unless otherwise indicated]: WA, BENTON [on Echinochloa 
crus-gallit, T. aestivum, & ZEA MAYs*|—-WSU-Prosser, 1-X-95 (95G668); nr Paterson, 12-X-95 (95G688): 
Cemetery Rd (3 mi east of Hwy 221), 12-X-95 (95G690); DouGLas—Waterville, 12-VIII-95 (95B016, 
95B017, 95B018); YAKIMA [*]. [Following collections on Zea mays]: AsoTIN—Chief Timothy HMU, 16- 
VIII-95 (95T439); BENTON [on Echinochloa crus-galli & Zea mays *|; BENTON—-WSU-Prosser, 24-VII- 
95, 18-IX-95, 6-X-95, 28-IX-96, 3-XI-97 (95G511, 95G601, 95G683, 95G684, 96G591, 97G518); Prosser, 
21-IX-96 (96G626); Grandview, 24-VIII-95 (95G513); GRANT—Ephrata, 8-VII-97 (97K058); WALLA 
WALLA—Hwy 124 (nr Ice Harbor Dam), 5-IX-95, 5,18-X-95 (95G543, 95G675, 95G699): Eureka, 5-X- 
95 (95G681, 95G698); Prescott, 30-X-97 on Phalaris arundinacea (97G516); YAKIMA—Grandview, 25- 
VIII-95, 18-IX-95, 24-X-95, (95G520, 95G602, 95G702); Harrah, 19-[X-95 (95G604); nr Toppenish, 19- 
IX-95, 16-X-95, 5-XI-95 (95G603, 95G695, 95G745). 

Schizaphis graminum (Rondani): WA, BENTON—15-VI-82, (82K001). 

Sitobion avenae (F) [on Triticum aestivum unless otherwise indicated]: WA, Asotin [*]; BENTON—-WSU- 
Prosser, 3,11,18 & 25-VII-97 (97F192, 97F193, 97F199, 97F200, 97F209, 97F215, 97F223, 
97F231,97F249, 97F252, 97F260, 97F263, 97F264, 97F265, 97F273), & 18-VII-97 on Setaria lutescens 
(97G243); KLickIrAtT—Bickleton Hwy & County Line Rd, 7-VII-97 (97G224). 

Sitobion sp.: WA, KLickirAt—Trout Lk, 11-VII-95 on Holodiscus discolor (95G340*#). 

Tuberculatus columbiae Richards: WA, YAKIMA—YIR, Mill Cr Guard Station, 6-VI-95 on Quercus garryana 
(95G112, 95G113), & on Salix sp. (95G114). 

Uroleucon cirsii (L.): WA, KLICKITAT—Brooks Memorial SP, 21-[X-95 on Cirsium sp. (95K162+); PIERCE [on 
Cirsium arvense*). 

Uroleucon sp.: ID, KOOTENAI—Rathdrum, 29-VII-95 (95G443}). 

Undetermined spp.: ID, LATAH—Moscow, 29-VIII-96 on Coriandrum sativum (96T513); BENTON—Prosser, 
1-VIII-95 on Portulaca oleracea (95A028); Kittiras—Stampede Pass, 8-VIII-96 on Epilobium 
angustifolium (96K146); LEwis—RNP, 8-IX-95 on Luetkea pectinata (95K115); WHITMAN [on Lupinus 
sp.*]—Pullman, 19-VI-96, 3-VII-96 on P. malus (96T042); YAKIMA—Grandview, 12-X-96 on Sonchus sp. 
(96G647). 


Lysiphlebus utahensis (Smith) 

Aphis canae Williams: WA, FRANKLIN—Kahlotus Hwy, 11-VI-96 on Artemisia tridentata (96G2427+). 

Artemisaphis artemisicola (Williams) [on Artemisia tridentata]: WA, FRANKLIN—Kahlotus Hwy, 28-V-96 & 
11-VI-96 on Artemisia tridentata (96G096, 96G242++); OKANOGAN—Bridgeport SP, 9-IX-97 (97K018); 
YAKIMA—Naught Rd, 18-VI-97 (97G1037#). 

Epameibaphis atricornis G&P: WA, BENTON—Byron Rd, 14-V-97 on Artemisia tridentata (97G024+#). 

Flabellomicrosiphum tridentatae (Wilson): WA, KLICKITAT—nr Roosevelt, 16-IV-96 on Artemisia tridentata 
(96G0127+#). 

Obtusicauda artemisiphila (Knowlton & Allen): WA, KLIicKkirAt—nr Roosevelt, 16-IV-96 on Artemisia 
tridentata (96GO012+#). 

Obtusicauda coweni (Hunter) [on Artemisia tridentata}: WA, ASoTIN—Asotin Cr HMU, 7-V-96 (96T 1427); 
YAKIMA [*]. 

Obtusicauda filifoliae (G&P) [on Artemisia tridentata): WA, BENTON—Byron Rd, 14-V-97 (97G0247#); 
KLICKITAT [*]—nr Roosevelt, 16-IV-96 (97G0127#). 

Obtusicauda sp.: WA, KLICKITAT [on Artemisia tridentata*}. 

Pseudoepameibaphis tridentatae (Wilson) [on Artemisia tridentata}: WA, BENTON—Byron Rd, 14-V-97 
(97G024++); KiLickiITAt—nr Roosevelt, 16-IV-96 (96G012+£); YAKImMA—Naught Rd, 18-VI-97 
(97G1037+#). 


Lysiphlebus sp. 
Aphis helianthi Monell: WA, PIERCE [on Heracleum lanatum*). 
Aphis sp.: WA, PIERCE [on Heracleum lanatum*]; YAKIMA [on unknown Umbelliferae*]. 
Brachycaudus tragopogonis (Kaltenbach): WA, KLICKITAT [on Tragopogon dubius*]. 


716 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Genus MONOCTONUS Haliday 
Monoctonus campbellianus Pike & Stary, new species 
(Figs. 13-24) 


Diagnosis.—The new species does not key to any of the North American species 
because of its reduced wing venation. According to the classification by van Achterberg 
(1989), it is part of the M. cerasi (Marshall) group. [Note, a part of the specimens of 
some species classified by Stary & Smith (1976) as European species, but distributed in 
N. America, are probably this new species]. 

Etymology.—The name of the new species is derived from the type locality, Campbell 
Lake, Skagit Co., Washington. 

Description.—Female. Head: Eyes with sparse setae in lower half. Tentorio-ocular line 
subequal to % of intertentorial line; malar space somewhat longer than /; of eye length, 
equal to half of intertentorial line. Antenna 13-segmented, short, as long as head, 
mesosoma and metasomal tergum | together, thickened to apex. Flagellomere 1 [F,] (Fig. 
15) length 3.5 width, without placodes, setae equal to segment diameter; F, (Fig. 16) 
slightly shorter than F,, about % wider than F,, with 3 placodes; preapical F-segments 
(Fig. 17) (dength/width = 1.7) % wider than F,. Mesosoma: Antescutal depression evident 
(Fig. 20). Mesonotum with notauli distinct anteriorly, smooth except for sparse setae 
tracing effaced notauli on the disc (Fig. 13). Propodeum (Fig. 14) with narrow central 
pentagonal areola. Forewing (Fig. 18): Stigma length about 3.5 width; distal abscissa 
of Rl (= metacarpus) short, about /% of stigma length; r + 3RS vein (radial sector) 
distinctly sclerotized (insertion point for completely effaced 2RS vein defines separation 
between r and 3RS veins). Stigma width 2 length of r-vein; 3RS vein 2—3X length of 
r-vein. Cubital cell narrow, distinctly complete or closed (Fig. 19). Legs: Femora (Fig. 
22). Metasoma: Metasomal tergum | (Fig. 21) length nearly 2 width at spiracles, disc 
coarsely rugose; spiracular tubercles positioned 1/3 from anterior margin; distance 
between spiracles and apex greater than width at spiracles. Genitalia (Fig. 23): Ovipositor 
sheath (Fig. 24) distinctly ploughshare-shaped. Coloration: Head dark brown; lower part 
of gena and mandibles yellow brown, palpi yellow. Antennae brown, except scape, pedicel 
and F, yellow. Mesosoma brown, sometimes propleurae, and part of mesopleurae and 
propodeum yellow brown. Wings subhyaline, venation light brown. Legs yellow, apices 
of tarsi darkened. Metasomal tergum | and central part of tergum 2 yellow to yellow 
brown, otherwise metasoma brown, the apex yellow brown to light brown. Ovipositor 
sheaths concolorous with the apex of metasoma, upper portion darker. Length of body: 
about 2.0—2.3 mm. 

Male.—Antenna 16-segmented. Head brown. Antennae brown, articulation between 
pedicel and F, lighter. Mandibles yellow brown, palpi yellow. Mesosoma brown. Legs 
yellow brown, coxae darkened. Metasoma brown, tergum 1 yellow brown. 

Holotype.—?, USA, WA, SKAGIT Co., Campbell Lake, 04-VI-1996, G. Graf & P. Stary 
collectors (collection code, 96G165). Host aphid (reared from a mixed collection): 
Brachycaudus helichrysi (Kaltenbach), Hyalopterus pruni (Geoffroy), and Phorodon 
humuli (Schrank) on Prunus sp. Deposited in USNM. 

Paratypes.—31 specimens, collection data same as holotype; deposited in part in 
collections of WSU-Prosser (16 2, 6 3) and P. Stary, Ceské Budejovice, Czech Republic 
COR 22s): 


VOLUME 102, NUMBER 3 Uy 


i 
insertion point 
of effaced 2RS 


Figs. 13-24. Monoctonus campbellianus, 2 [illustrations not to equal scale]. 13, Mesonotum. 14, Propo- 
deum. 15—17, Flagellomeres 1, 2, and preapical. 18, Fore wing, in part. 19, Cubital cell. 20, Mesosoma— 
pronotum with antescutal depression. 21, Metasomal tergum 1. 22, Femora. 23, Genitalia. 24, Ovipositor sheath. 
Abbreviations: 2RS, 3RS, and r, wing veins. 


718 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Monoctonus pacificus Pike and Stary, new species 
(Figs. 25—32) 


Diagnosis.—The new species is similar to M. nervosus (Haliday) (see van Achterberg 
1989), but differs from the latter in coloration, number of antennal segments, absence of 
placodes on flagellar segments 1—4, and arcuate r-vein in the forewing. 

Etymology.—The name of the new species is derived from the general type locality, 
the Pacific Northwest, USA. 

Description.—Female. Head: Eyes with sparse setae in lower half. Tentorio-ocular line 
subequal to 4 of intertentorial line; malar space 2 length of intertentorial line or /; of 
eye length. Antenna 17—18-segmented, filiform, reaching to half of metasoma. 
Flagellomere 1 [F,] (Fig. 27) length 4.0 width, the setae as long to longer than segment 
diameter. F, (Fig. 28) subequal to F,. F,_, without placodes; F,, 1 placode; F, 3 placodes. 
Preapical F-segments (Fig. 29) about /, wider than F,. Mesosoma: Antescutal depression 
evident (Fig. 32). Mesonotum with large bare areas in central and lateral lobes. Propodeum 
(Fig. 25) with narrow central areola. FOREWING (Fig. 31): Stigma length 6.0 width; distal 
abscissa of RI (metacarpus) short, equal to ¥% of stigma length; r + 3RS vein (radial 
sector) distinctly sclerotized; r-vein arcuate, slightly shorter than stigma width; 3RS equal 
to r-vein. 2RS vein colorless but distinct. RS + M vein distinct. Metasoma: Metasomal 
tergum | (Fig. 26) length 2X width at spiracles, with spiracles positioned % from anterior 
margin; disc coarsely rugose. Genitalia (Fig. 30): Ovipositor sheath ploughshare-shaped. 
Coloration: Largely yellow. Head, including mandibles brown; palpi yellow; antennae 
light brown, except scape, pedicel, F,_, yellow (sometimes F; and F, infuscated). Length 
of body: about 2.3 mm. 

Male. Antenna 19—20-segmented. Coloration similar to female, metasoma somewhat 
darkened. 

Holotype.—?, USA, WA, YAKIMA Co., nr Clear Lake, 15-VI-1996, K. S. Pike collector, 
(collection code, 96K099). Host aphid: Macrosiphum tuberculaceps (Essig) on Achlys 
triphylla. Deposited in USNM. 

Paratypes.—5 specimens, USA, WA, YAKIMA Co., N Fork of Tieton River, 27-VI-1996. 
Host aphid: Macrosiphum tuberculaceps (Essig) on Achlys triphylla (96G298), deposited 
in part in collections of WSU-Prosser (3 2, 1 6) and P. Stary, Ceské Budejovice, Czech 
Republic (1 @). 

Other material reared from: 

Macrosiphum tuberculaceps (Essig): WA, YAKIMA, Morse Cr, 24-VII-97 on Achlys 
triphylla (97G255), N Fork of Tieton River, 27-VI-1996 on Achlys triphylla (96G298); 
nr Clear Lake, 15-VI-1996, on Achlys triphylla (96K099). 


Monoctonus washingtonensis Pike & Stary 
Aphis nr oenotherae Oestlund: WA, GARFIELD—Spruce Springs, 22-VIII-95 on Epilobium sp. (95T4627). 
Diuraphis noxia (Kurdjumov): WA, BENTON [on Triticum aestivum*]. 
Rhopalosiphum padi (L.): WA, BENTON [on Triticum aestivum*]. 


Monoctonus spp. 
Acyrthosiphon pisum (Harris): WA, KittiTAs [on Rhododendron albiflorum*]. 
Illinoia rhododendri (Wilson): WA, KittiTas [on Rhododendron albiflorum* ]. 
Illinoia sp.: WA, YAKIMA [on Rhododendron albiflorum*]. 
Macrosiphum euphorbiae (Thomas): WA, SKAMANIA—Twin Falls CG, 4-VIII-95 on Pteridium aquilinum 
(95K099). 
Macrosiphum walkeri Robinson: OR, BENTON—McDonald State Forest, 21-V-91 on Polypodium sp. (91JO03). 
Metopolophium dirhodum (Walker): WA, Kirtiras [on Rhododendron albiflorum*}. 


VOLUME 102, NUMBER 3 TS 


BY) 


Figs. 25-32. Monoctonus pacificus, 2 [illustrations not to equal scale]. 25, Propodeum. 26, Metasomal 
tergum 1. 27-29, Flagellomeres 1, 2, and preapical. 30, Genitalia. 31, Fore wing, in part. 32, Mesosoma— 
pronotum with antescutal depression. Abbreviations: R1, distal abscissa of postmarginal vein [= metacarpus]; 
2RS, 3RS, RS+M, and r, wing veins. 


720 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Genus PAUESIA Quilis 


Pauesia ahtanumensis Pike & Stary 
Cinara ponderosae (Williams) [on Pinus ponderosa unless otherwise indicated]: ID, LATAH—Moscow Mtn, 
6-VII-96 on Pinus sp. (961195); MT, LincoLN—Libby, 16-VI-95 (95G171); WA, AsotiIn—De Spain 
Springs, 5-VII-96 (96T149); Kiickirat—Bickleton, 18-VI-97 (97G112); OKANOGAN—Carlton, 11-IX-97 
(97K034); SKAMANIA & YAKIMA [*]. 
Cinara sp.: WA, AsoTiIN—Copper Canyon, 18-VII-96 on Pinus ponderosa (961308). 


Pauesia bicolor (Ashmead) 
Cinara sp.: ID, BouNDARY—Bonners Ferry, 29-VII-95 on Pinus contorta (95G444). 


Pauesia juniperaphidis (Gahan) 
Cinara pilicornis (Hartig): WA, YAKIMA [on Picea pungens*}. 


Pauesia pahtonis Pike & Stary 
Cinara ponderosae (Williams): WA, SKAMANIA [on Pinus ponderosa*}. 


Pauesia ponderosaecola Pike & Stary 
Cinara ponderosae (Williams) [on Pinus ponderosa unless otherwise indicated]: MT, LINCOLN—Libby, 16- 
VI-95 (95G171); WA, AsoTin—De Spain Spring, 5-VII-96 (96T149, 96T151); Krirtiras—S Fork 
Manastash Cr, 25-VI-97 (97G136); KLICKITAT [*]—Bickleton, 18-VI-97 (97G112); WHITMAN—Kamiak 
Butte, 7-VII-96 on Pinus sp. (961203); YAKIMA [*]—YIR, White Deer Cr, 11-VII-95 (95K050). 


Pauesia pseudotsugae Pike & Stary 
Cinara pseudotaxifoliae Palmer: WA, Kittitas [on Pseudotsuga menziesii*]. 


Pauesia spp. 
Cinara brevispinosa (G&P): WA, SPOKANE—Chapman Lk, 1-VI-95 on Pinus ponderosa (95T046B); YAKIMA 
[on Pseudotsuga menziesii*|—YIR, Potato Hill Guard Station, 11-VII-95 on Pinus contorta (95K063). 
Cinara ponderosae (Williams): WA, SPOKANE—Deer Pk, 23-V-95 on Pinus ponderosa (95KO15). 
Cinara pseudotsugae (Wilson): WA, YAKIMA [on Pseudotsugae menziesii*). 
Cinara sp.: WA, YAKIMA—N Fork Tieton River, 27-VI-96 on Abies amabilis (96G293). 
Undetermined sp.: WA, YAKIMA—Clear Lk, 15-VI-96 on Pinus ponderosa (96K087). 


Genus PRAON Haliday 


Praon americanum (Ashmead) 
Aphis coweni Palmer: WA, Kittrras—Stampede Pass, 8-VIII-96 on Veratrum viride (96K147). 
Aphis salicariae Koch: WA, YAKIMA—YIR, nr. Mt. Adams Lk., 19-IX-95 on Epilobium angustifolium 
(95G6107). 


Praon artemisaphis Smith 
Obtusicauda coweni (Hunter): WA, YAKIMA [on Artemisia tridentata*). 
Obtusicauda sp.: WA, KLICKITATt & YAKIMA [on Artemisia tridentata *]. 


Praon artemisicola Pike & Stary 
Artemisaphis artemisicola (Williams): WA, BENTON—Six Prong Rd, 24-IV-97 on Artemisia tridentata 
(97G0087+). 
Epameibaphis atricornis G&P [on Artemisia tridentata]: WA, KLIckitaT [*+], & YAKIMA [esta 
Flabellomicrosiphum sp. [on Artemisia tridentata]: WA, KLICKITAT [**+], & YAKIMA [ale 
Pleotrichophorus quadritrichus (Knowlton & Smith): WA, YAKIMA—Mabton, 24-IV-97 on Artemisia 
tridentata (97G0077). 


Pleotrichophorus sp.: WA, KLickirat—Alderdale Rd, 9-V-95 on Artemisia tridentata (95GO287¢). 
Pseudoepameibaphis tridentata (Wilson) [on Artemisia tridentata): WA, BENTON [*], KLICKITAT [*+]J— 
Alderdale Rd, 9-V-95 (95G0287); YAKIMA [*]. 


Praon coniforme Pike and Stary, new species 
(Figs. 33-41) 


Diagnosis.—The new species belongs to a species group characterized by the absence 
of an m-cu vein in the forewing. Other distinguishing characters are antenna 16-17 
segmented (in female), flagellomere 1 (F,) yellow, and ovipositor sheath coniform in 


VOLUME 102, NUMBER 3 721 


Ni & 

SSN Nerval 
\ 

to NaS 

7 


/ 
7 
- 


36 


37 


Figs. 33-41. Praon coniforme, @ [illustrations not to equal scale]. 33, Mesonotum. 34, Propodeum. 35, 
Metasomal tergum 1. 36—38, Flagellomeres 1, 2, and preapical. 39, Fore wing, in part. 40, Genitalia. 41, Ovi- 
positor sheath. Abbreviations: R1, distal abscissa of postmarginal vein [= metacarpus]; r+2RS, wing vein. 


shape. The new species is similar to Praon yakimanum (see Pike & Stary 1995), but 
differs from the latter in antennal segment number (P. yakimanum, 15-segmented [rarely 
16]), shape of the ovipositor sheath, and aphid host. At present, the new species is only 
known from snowberry aphid, Aphthargelia symphoricarpi (Thomas). 

Etymology.—The name of the new species is derived from the shape of the ovipositor 
sheath that resembles a cone. 

Description.—Female. Head: Eyes medium sized, with sparse setae in lower half. 
Tentorio-ocular line subequal to 1/4 of intertentorial line; malar space 1.5x< length of 
tentorio-ocular line or slightly longer than ¥, of eye length. Antenna 16—17 segmented, 
filiform, slightly thickened in apical third, reaching to half of metasoma. Flagellomere 1 
[F,] (Fig. 36) slightly more than 4.0 width; 1.6 as long as F,; setae slightly longer 
than segment diameter; without placodes. F, (Fig. 37) width equal to F,, with 1 placode. 
Preapical F-segments (Fig. 38) 1.3 width of F,. Mesosoma: Mesonotum (Fig. 33) with 
bare areas on the lateral lobes. Propodeum (Fig. 34) pubescent. FOREWING (Fig. 39): 
Stigma length 3.5 width; distal abscissa of R1 (metacarpus) longer than half stigma 
length; r + 3RS vein (radial sector) slightly longer than distal abscissa of R1; m-cu vein 
absent. Metasoma: Metasomal tergum 1 (Fig. 35) somewhat longer than width at spiracles 
(7:6); distance between spiracles and anterior margin %4 shorter than width at spiracles. 
Genitalia (Fig. 40): Ovipositor sheath (Fig. 41) cone-like in shape. Coloration: Head 
dark brown, mouthparts bright yellow. Antenna brown; apex of pedicel and F, yellow. 
Mesosoma dark brown. Wings subhyaline, venation light brown. Legs yellow, apices of 


22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


tarsi infuscate. Metasomal tergum | yellow brown, darker in basal portion. Metasoma 
brown. Ovipositor sheaths dark brown. Length of body: about 2.4 mm. 

Male.—Antenna 19—20-segmented, filiform, longer than the body. Antenna brown, F, 
with a narrow yellow base. Legs light brown. 

Holotype.-—?, USA, WA, YAKIMA Co., Maloy Rd and Wenas Creek, 30-V-1996, G. 
Graf collector, (collection code, 96G127). Host aphid: Aphthargelia symphoricarpi 
(Thomas) on Symphoricarpos sp. Deposited in USNM. 

Paratypes.—3 specimens, collection data same as holotype, deposited in part in 
collections of WSU-Prosser (2 6) and P. Stary, Ceské Budejovice, Czech Republic (1 2). 

Other Material reared from: 

Aphthargelia symphoricarpi (Thomas) [on Symphoricarpos sp. unless otherwise 
indicated]: WA, KittirAs—Lost Lake, 29-VI-1994 (94G005) & 6-VIII-1997 (97G334); 
KLICKITAT—Trout Lake, 28-VI-1997 (97G159); SPOKANE—Cooper Rd, 29-VI-95 
(95T178); Fish Lake, 14-VI-1997 on S. occidentalis (97KO04); YAKIMA—Maloy Rd & 
Wenas Cr., 11-VI-1997 (97G066). 


Praon exsoletum (Nees) 

Therioaphis riehmi (Borner) [on Melilotus sp. unless otherwise indicated]: WA, BENTON—-WSU-Prosser, | 1- 
VII-97 on M. officinalis (97F218+); YAKIMA—Grandview, 22-VI-96. (96G261); nr Parker, 29-I[X-95 
(95G665+); YIR, Signal Peak, 19-IX-95 (95G6257). 

Therioaphis trifolii (Monell) [on Medicago sativa): WA, BENTON [*]; YAKIMA, nr Union Gap, 23-X-95 
(95G7217). 

Therioaphis sp.: WA, BENTON—-WSU-Prosser, 28-VI-95 on Medicago sativa (95G2777). 


Praon fulvum Pike and Stary, new species 
(Figs. 42—50) 


Diagnosis.—Yellow coloration of F, (flagellomere 1) and F, keys the new species to 
P. gallicum Stary (see key by Johnson 1987), but it differs from the latter in the number 
of antennal segments [P. fulvum with 18—19 (rarely 20) vs. P. gallicum with 15—16 (rarely 
17)], the presence of the m-cu vein in the forewing (absent with P. gallicum), and the 
mesonotum pubescence on the lateral lobes of the mesonotum (large bare areas with P. 
gallicum). 

Etymology.—The name of the new species is derived from its prevailing body 
coloration. 

Description.—Female. Head: Eyes medium sized, with sparse setae in lower half. 
Tentorio-ocular line % of intertentorial line or 4% of malar space. Antenna 18-19 (rarely 
20) segmented, filiform, slightly thickened towards apex, subequal to body length. 
Flagellomere 1 [F,] (Fig. 45) long, length nearly 5X width, setae slightly longer than 
segment diameter, without placodes. F, (Fig. 46) % shorter than F,, without placodes. 
Preapical F-segments (Fig. 47) 1/8 wider than F,. Mesosoma: Mesonotum (Fig. 42) with 
very small to no areas on the lateral lobes without setae. Propodeum (Fig. 49) pubescent, 
somewhat bare in center. FOREWING (Fig. 48): Stigma length 3.5 width; distal abscissa 
of R1 (metacarpus) slightly longer than half stigma length; RS + M vein distinct, 
subcolored in basal 4—%, remaining portion colorless; m-cu vein complete, subcolored. 
Metasoma: Metasomal tergum 1 (Fig. 50) longitudinal, length %4 longer than width at 
spiracles; width at spiracles Y; longer than the distance from spiracles to anterior margin. 
Genitalia (Fig. 43): Ovipositor sheath (Fig. 44) long, subarcuate. Coloration: Body 
almost completely yellow. Head yellow, frons more or less brown. Apices of mandibles 
brown. Antenna brown; scape, pedicel, F, and F, yellow. F, sometimes infuscated at apex. 


VOLUME 102, NUMBER 3 123 


Figs. 42-50. Praon fulvum, @ [illustrations not to equal scale]. 42, Mesonotum. 43, Genitalia. 44, Ovipositor 
sheath. 45—47, Flagellomeres 1, 2, and preapical. 48, Fore wing, in part. 49, Propodeum. 50, Metasomal tergum 
1. Abbreviations: R1, distal abscissa of postmarginal vein [= metacarpus]; RS+M and M-cu, wing vein. 


Mesosoma yellow. Wings subhyaline, venation light brown. Legs yellow. Metasoma 
yellow. Ovipositor sheaths light brown. Length of body: about 2.0—2.6 mm. 
Male.—Unknown. 
Holotype.— ?, USA, WA, Lewis Co., Mt. Rainier Nat. Park, Stevens Canyon, 23-VII- 


724 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


1996, G. Graf collector, (collection code, 96G357). Host aphid: Macrosiphum euphorbiae 
(Thomas) on Crepis sp. Deposited in USNM. 

Paratypes.—6 specimens, collection data same as holotype, deposited in part in 
collections of WSU-Prosser (3 ¢) and P. Stary, Ceské Budejovice, Czech Republic (3 2). 

Other material reared from: 

Macrosiphum euphorbiae (Thomas): WA, KLICckIrAt—Cleveland, 7-VII-1997, on 
Lupinus sp. (97G230). 

Macrosiphum tuberculaceps (Essig): WA, YAKIMA—Wenas Creek & Maloy Rd, 11-VI- 
1997 on Achlys triphylla, (96G298). 

Macrosiphum sp.: WA, YAKIMA—Wenas Creek & Maloy Rd, 11-VI-1997 on Spiraea 
pyramidata (97G073). 


Praon gallicum Stary 

Diuraphis noxia (Kurdjumoy) [on Triticum aestivum]: WA, BENTON—WSU-Prosser, 22-VII-97 (97F295); 
KLickIrAt—Crider Valley Rd, 22-VII-97 (97G250++); YAkIMA—nr Sunnyside, 17-VII-97 (97G2467). 

Metopolophium dirhodum (Walker) [on Triticum aestivum]: WA, YAKIMA—Glade Rd, 7-VII-97 (97G2177); 
nr Sunnyside, 17-VII-97 (97G2467). 

Rhopalosiphum padi (L.): WA, GRANT—Ephrata, 8-VII-97 on Zea mays (97K058); KLICKITAT—Crider Valley 
Rd, 22-VII-97 on Triticum aestivum (97G2507¢). 

Sitobion avenae (FE) [on Triticum aestivum]: WA, KLIcKiIrat—Crider Valley Rd, 22-VII-97 (97G2507+); 
Y AKIMA—Glade Rd, 7-VII-97 (97G2177). 


Praon humulaphidis Ashmead 
Acyrthosiphon macrosiphum (Wilson): MT, LINCOLN—Libby, 6-VII-96 on Amelanchier alnifolia (96G322). 
Macrosiphum euphorbiae (Thomas): WA, WHITMAN—Pullman, 18-VI-96 on cultivated Rosa sp. (961037) & 
5-VIH-96 on unknown plant (96T176). 
Macrosiphum sp. 1D, BENEWAH—McCroskey SP, 7-VI-96 on native Rosa sp. (96TO005). 
Nasonovia ribisnigri (Mosley): WA, WHITMAN—Pullman, 5-VIU-96 on unknown plant (96T177). 
Illinoia sp.: 1D, SHOSHONE—Thomas Pass, 29-VII-95 on Vaccinium sp. (95G472). 
Illinoia sp.?: WA, CHELAN—Chatter Cr CG, 1-VIII-95 on Aruncus sylvester (95G487). 


Praon occidentale Baker 

Acyrthosiphon lactucae (Passerini) [on Lactuca serriola}: ID, BENEWAH—McCroskey SP, 31-VIII-95 
(95T514+); WA, BENTON—WSU-Prosser, 11,22-V-95 (95G034, 95G081); Prosser, 22-VI-97 (97G121); 
WHITMAN—Chambers, 19-IX-95 (95T581); Nisqually John Landing, 19-IX-95 (95T584); Pullman, 2-VIII- 
96 (961386); YAKIMA—Grandview, 23-VI-96 (96G263); Wenas Lk, 11-VI-97 (97G055). 

Acyrthosiphon pisum (Harris): WA, BENTON—28-VI-95 on Medicago sativa (95G277+); Prosser, 23-VII-96 
on Trifolium repens (96G393); GRANT—WSU-Othello, 26-VI-95 on Medicago sativa (95B008); 
SKAMANIA—Underwood, 11-VII-95 on Lathyrus sp. (95K072). 

Amphorophora geranii G&P: WA, WHITMAN—Long Rd nr Hole in the Ground, 18-V-95 on Geranium sp. 
(95T0127). 

Aphis craccivora Koch: ID, LAraH—Moscow Mtn, 3-VIII-96 on Trifolium sp. (96T4007+). 

Aphis helianthi Monell: WA, GARFIELD—Stentz Springs, 22-VIII-95 on Osmorhiza chilensis (95T452). 

Aphis lugentis Williams: WA, YAKIMA—Bird Cr Meadows, 8-[X-94 on Arnica sp. (94R172). 

Brachycaudus helichrysi (Kaltenbach): WA, GARFIELD—FR160 & FR40, 21-I[X-95 on Anaphalis margaritacea 
(95T5957+£); PIERCE—RNP, Tipsoo Lk, 5-IX-97 on Aster sp. (97G364++£); YAKIMA—Mabton-Bickleton Rd, 
Glade Cr, 7-VII-97 on Artemisia ludoviciana (97G219). 

Capitophorus elaeagni (del Guercio): WA, YAKIMA—Grandview, 28-VI-96 on Cirsium vulgare (96G3097+). 

Chaetosiphon thomasi HRL: WA, PIERCE—WSU-Puyallup, 10-VI-96 on cultivated Fragaria sp. (96T022). 

Diuraphis noxia (Kurdjumov) [on Triticum aestivum]: WA, BENTON [*]; KLICKIrAT—Bickleton, 29-VII-97 
(97G300). 

Dysaphis plantaginea (Passerini): WA, YAKIMA—Grandview, 10-VI-96 on Pyrus malus (96G214). 

Ericaphis wakibae (Hottes): WA, YAKIMA—N Fork Tieton River, 15-VI-96 on native Rosa sp. (96KO0907+). 

Hyperomyzus nigricornis (Knowlton) [on Hieracium sp. unless otherwise indicated]: ID, BENEWAH— 
McCroskey SP, 31-VII-95 (95T514+); LATAH—Moscow Mtn, 15-VIII-96 on Compositae (961446); MT, 
SANDERS—nr Thompson Pass, 29-VII-95 (95G462+); WA, GARFIELD—Alder Thicket, 21-[X-95 (95T587); 
Kitriras—Buck Meadows, 16-VIII-96 (96G493); Manastash Rd, 25-VI-96 (96G415); KLICKITAT— 


VOLUME 102, NUMBER 3 725 


Cleveland, 18-VI-96 on Madia minima (96K109); SPoKANE—Mt Spokane Hwy (mi marker 18.5), 22-VIII- 
96 on Compositae (96T487, 96T488); YAKIMA—Bird Cr Meadows, 26-VI-95 (95A0167). 

Hyperomyzus sp.: WA, Kirriras—S Fork Manastash Cr, 24-VII-95 on Hieracium albiflorum (95G388): 
SPOKANE—nr Peone, 24-VIII-95 on Centaurea pratensis (95T4737+); WHITMAN—Pullman, 10-VIII-95 on 
Lapsana communis (9574364). 

Illinoia davidsoni (Mason): MT. SANDERS—nr Thompson Pass, 29-VII-95 on Arnica sp. (95G463); WA, 
KitTTITAs—Quartz Mtn, 25-VII-96 on Arnica cordifolia (96G406++). 

Illinoia richardsi: WA, GARFIELD—FR160 & FR40, 21-IX-95 on Anaphalis margaritacea (95T595+%). 

Illinoia nr thalictri (MacGillivray): WA, GARFIELD—Stentz Springs (FR 40), 22-VIII-95 on Thalictrum sp. 
(95T454). 

Ilinoia spp.: ID, BENEWAH—McCroskey SP 7-VI-96 on unknown plant (96T006); WA, GarFIELD—Stentz 
Springs, 1-VIIH-95 on Aquilegia formosa (95T362+): KITTITAS [on Aquilegia formosat*]; PlERCE—RNP. 
Tipsoo Lk, 11-IX-96 on Carex sp. (96G5427 +). 

Macrosiphoniella ludovicianae (Oestlund): WA, Asotin—Asotin Cr WLA, 26-VI-96 on Artemisia ludoviciana 
(96T048). 

Macrosiphoniella nr ludovicianae (Oestlund):; WA, YAKIMA [on Artemisia ludoviciana*]. 

Macrosiphum albifrons Essig: WA, PIERCE—RNP. Tipsoo Lk, 11-[X-96 on Lupinus sp. (96G547). 

Macrosiphum euphorbiae (Thomas): ID, LATAH—Moscow Mtn, 6-VII-96 on Castilleja sp. (96T188); 6-VII- 
96, on native Rosa sp. (96T184); 6-VII-96 on Compositae (96T185, 96T192): MT, LINCOLN—Libby, 15- 
VI-95 on native Rosa sp. (95G168); SANDERS—nr Thompson Pass, 29-VII-95 on Hieracium sp. (95G4627); 
WA, AsoTIN—De Spain Springs, 1-VIII-95 on Senecio sp. (951381), 26-VI-96 on unknown plant (96T0S51), 
5-VII-96 on Compositae (96T159), 11-VII-96 on Potentilla sp. (96T250) & Compositae (96T251); 
BENTON—WSU-Prosser, 27-X-95 on Chenopodium album (95K2187}), & 27-X-95 on cultivated Rosa sp. 
(95K220+); Kirtrras—Buck Meadows, 9-VII-96 on Agoseris elata (96G341), 9-VII-96 on native Rosa Sp. 
(96G335); Manastash Cr, FR125, 16-VIII-96 on Silene sp. (96G497): Manastash Ridge, 2-IX-95 on 
Gnaphalium microcephalum (95G537): Lost Lk Trail, 6-VIH-97 on Luina nardosmia (97G327); Quartz 
Mtn, 25-VII-96 on Arnica cordifolia (96G406+); PEND OREILLE, Tiger Meadows, 9-VII-96 on Compositae 
(96T221); SPOKANE—Mt Spokane SP, 24-VIII-95 on native Rosa sp. (95T491+); STEVENS—Little Pend 
Oreille WLA, 9-VII-96 on Compositae (96T210); WHITMAN—Pullman, 10-VIII-95 on Lapsana communis 
(95T436+) & 18-VI-96 on cultivated Rosa sp.(96T037); WA, YaKIMA—Bird Cr Meadows, 26-VI-95 on 
Hieracium sp. (95A016+); Grandview, 28-VI-96 on Cirsium vulgare (96G309+); Hwy 410 & Sawmill Flat, 
28-VII-97 on native Rosa sp. (97G2897). 

Macrosiphum osmaroniae (Wilson): WA, WHATCOM—Bellingham, 3-V-96 on Oemleria cerasiformis 
(96K013); 

Macrosiphum rosae (L.) [on cultivated Rosa sp.]: WA, BENTON—-WSU-Prosser, 27-X-95 (95K2207); nr 
Prosser, Richards Rd, 22-VI-97 (97G122); YAKIMA [+*]—Grandview, 13 & 21-V-97 (97G021++4, 97G037). 

Macrosiphum sp.: ID, BENEWAH—McCroskey SP. 7-VI-96 on native Rosa sp. (96T005); WA, AsotTin—nr 
Anatone, 24-VIII-93 on Triticum aestivum (93T015); YAKIMA—nr Chinook Pass, Morse Cr, 24-VII-97 on 
Oplopanax horridum (97G256); Sunnyside, 24-VIII-95 on Zea mays (95G5197). 

Macrosiphum sp.?: WA, Kittiras—Manastash Ridge, 2 mi E of Lost Lk, 2-IX-95 on Silene sp. (95G545). 

Metopolophium dirhodum (Walker): WA, YAKIMA—N Fork Tieton River, 15-VI-96 on native Rosa sp. 
(96K090+F4). 

Metopolophium sp.?: WA, YAKIMA—YIR, Windy Point, 19-IX-95 on Poa sp. (95G634). 

Microlophium nr sibiricum: WA, YAKIMA—Sunnyside, 28-VI-95 on Urtica dioica (95G255). 

Myzus persicae (Sulzer): WA, BENTON—WSU-Prosser, 27-X-95 & 1-X-97 on Chenopodium album (95K218}+, 
97G476). 

Nasonovia aquilegiae (Essig): WA, GARFIELD—Stentz Springs, 1-VIII-95 on Aquilegia formosa (95T3627). 

Nasonovia polemonii (G&P): WA, KitrirAs—Quartz Mtn, 25-VII-96 on Polemonium occidentale (96G409). 

Nasonovia (Kakimia) sp.: WA, WHITMAN—Pullman, 10-VIII-95 on Lapsana communis (95743674). 

Nearctaphis bakeri (Cowen): ID, LATAH—-Moscow Mtn, 3-VIII-96 on Trifolium sp. (96T4007 +). 

Nearctaphis kachena (Hottes): WA, SPOKANE—nr Mt Spokane SP, 24-VIII-95 on Castilleja (95T485). 

Phorodon humuli (Schrank): WA, BENTON—WSU-Prosser, 9-IX-95 on Humulus lupulus (95K154). 

Rhopalosiphum insertum (Walker): WA, DouGLas—Orondo, 28-V-96 on Pyrus malus (96RO004). 

Rhopalosiphum padi (L.): WA, BENTON [on Triticum aestivum*]; GRANT—Ephrata 8-VII-97 on Zea mays 
(97K058); YAKIMA—Sunnyside, 24-VIII-95 on Zea mays (95G5197). 

Sitobion avenae (FE): WA, BENTON—WSU-Prosser, 3 & 11-VII-97 on Triticum aestivum (97F192, 97F211); 
Kittiras—Buck Meadows, 9-VII-96 on Dactylis glomerata (96G346); PiERCE—RNP, Tipsoo Lk, 11-IX- 
96 on Carex sp. (96G5427+#). 


726 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Uroleucon erigeronense (Thomas): WA, Kittiras—Lost Lk, 6-VIII-97 on Aster sp. (97G346); LEwis— 
PIERCE—RNP, Tipsoo Lk, 8-IX-95 & 5-IX-97 on Aster sp. (95K106, 97G3647). 

Uroleucon sonchi (L.): WA, FRANKLIN—Mesa, 8-IX-97 on Lactuca serriola (97KO06). 

Uroleucon zymozionense (Knowlton): WA, CHELAN—Swauk Pass, 1-VII-95, on Aster sp. (95G491). 

Uroleucon sp.: ID, BENEWAH—McCroskey SP, 31-VIII-95 on Compositae (95T528); WA, PEND OREILLE— 
Tiger Meadows, 9-VII-96 on Achillea millefolium (961222); PIERCE—RNP, Tipsoo Lk, 5-IX-97 on 
Anaphalis margaritacea (97G367); SKAMANIA [on Aster sp.*]; SPOKANE—nr Peone, 24-VIII-95 on 
Centaurea pratensis (95T4737*). 

Wahlgreniella nervata (Gillette): WA, BENToN—Grandview, 13-V-97 on cultivated Rosa sp. (97G0217++); 
YAKIMA—Hwy 410 & Sawmill Flat, 28-VII-97 on native Rosa sp. (97G2897$). 

Undetermined spp.: WA, BENTON—Prosser, 23-VII-96 on Sonchus oleraceus (96G389); KittiIras—S Fork 
Manastash Cr, 24-VII-95 on Potentilla sp. (95G395); WHITMAN—Pullman, 5-VII-96, on Geranium 
viscosissimum (96T171). 


Praon nr occidentale Baker 
Illinoia sp.: WA, YAKIMA—Blowout Mtn, end of Little Naches River Rd, 9-IX-97 on Rhododendron albiflorum 
(97G400). 
Uroleucon erigeronense (Thomas): WA, Kittiras—Lost Lk, 6-VIII-97 on Haplopappus hirtus (97G3407). 
Uroleucon nr ivae Robinson: WA, YAKIMA—Harrah, 29-IX-97 on Iva xanthifolia (97G465). 
Uroleucon suzannae Robinson: WA, Kirtrras—Lost Lk, 6-VIII-97 on Haplopappus hirtus (97G3407 +). 


Praon pequodorum Viereck 

Acyrthosiphon macrosiphum (Wilson): WA, AsoTin—nr Anatone, 18-VII-95, on Amelanchier alnifolia 
(95T266). 

Acyrthosiphon pisum (Harris) [on Medicago sativa unless otherwise indicated]: WA, ADAMS—WSU-Othello, 
26-VI-95 (95B008); BENTON [+*]—-WSU-Prosser, 27-VI-96 on Vicia angustifolia (Q96A009), 30-VI-97, & 
11-VII-97 (97F186, 97F202); GRANT—WSU-Royal City, 26-VI-95 (95B003, 95B006); Mattawa, 26-VI- 
95 (95B005); YAKIMA—Wenas Valley, 5 mi N of Selah, 30-V-96 on Melilotus sp. (96G109). 

Amphorophora rubi (Kaltenbach): WA, KLickirat—Trout Lk, 29-VI-97 on Rubus sp. (97G168). 

Illinoia sp.: WA, SPOKANE—Mt Spokane SP, 20-VII-95 on Vaccinium sp. (95T297). 

Therioaphis riehmi (Borner): WA, BENTON—-WSU-Prosser, 18 & 25-VII-97, on Melilotus officinalis 
(97F2547 4). 

Uroleucon ivae Robinson [on Iva xanthifolia]): WA, YAKIMA [*]—Grandview, 17,22, 30-VH-95 (95G313, 
95G349, 95G366). 

Uroleucon sonchi (L.): WA, YAKIMA—Grandview, 17-VII-95 on Sonchus oleraceus (95G348). 


Praon simulans (Provancher) 
Macrosiphum rhamni (Clarke) [det. by A. Jensen]: WA, Kinc—Stampede Pass, 8-VIII-96 on Pteridium 
aquilinum (96G441). 


Praon unicum Smith 

Acyrthosiphon lactucae (Passerini) [on Lactuca serriola}: WA, AsoTIN—Asotin Cr WLA, 7-IX-95 (95T547); 
BENTON—Prosser, 22-V-95 (95G079); CHELAN—Chelan SP, 11-[X-97 (97K038); KitTiTas [*]. 

Aphis coweni Palmer [on Veratrum viride]: WA, SKAMANIA [*]; YAKIMA [*]—Pleasant Valley CG, 24-VII-97 
(97G266). 

Aphis craccivora Koch: WA, WHITMAN—Pullman, 7-VII-96, 2-VIII-96 on Robinia pseudo-acacia (961199, 
96T393). 

Aphis nr fabae Scopoli: WA, YAKIMA—YIR, Windy Point, 19-I[X-95 on Cimicifuga laciniata (95G630). 

Aphis helianthi Monell: WA, CHELAN—Stevens Pass, 16-[X-95 on Epilobium angustifolium (95K1347¢); 
Kittiras—Lost Lk, 24-VII-95 on E. angustifolium (95G399+4); S. Fork Manastash Cr, 24-VII-95 on 
Heracleum lanatum (95G378); WHITMAN—Pullman, 1-VIII-95 on Coriandrum sativum (9513867); 
Y AKIMA—Ame rican River, 5-[X-97 on Heracleum lanatum (97G382); nr Wenas Lk, 17-VI-97 on Lomatium 
sp. (97G095+); YIR, Piscoe Cr, 25-VII-95 on Ligusticum grayi (95G432) & Osmorhiza occidentalis 
(95G428). 

Aphis nigratibialis Robinson: WA, Kirtrras—S. Fork Manastash Cr, mi marker 12, 25-VI-97 on Cornus 
stolonifera (97G129). 

Aphis oenotherae (Oestlund): ID, BENEWAH—McCroskey SP, 31-VII-95 on Epilobium angustifolium 
(95T533). 

Aphis nr oenotherae (Oestlund) [on Epilobium sp. unless otherwise indicated]: WA, BENTON—Kennewick, 
28-IX-96 (96G597); GARFIELD—FR 160 & FR 40, 21-IX-95 (95T594); Spruce Springs, 22-VIII-95 
(9514627); Stentz Springs, 14-[X-95 (95T565); YAKIMA—American River, 5-I[X-97 on Epilobium minutum 


VOLUME 102, NUMBER 3 727 


(97G375+); Pleasant Valley CG, 24-VII-97 on Epilobium minutum (97G267+); Wenas Cr, 13-VIII-96 
(96G467). 

Aphis pomi de Geer: WA, DouGLAs—Bridgeport, 17-VI-96 on Pyrus malus (96X007+#). 

Aphis rumicis L.: WA, BENTON, on [Rumex crispus*]. 

Aphis salicariae Koch [on Epilobium angustifolium]: ID, KOOTENAI—Rathdrum, 29-VII-95 (95G443+); WA, 
CHELAN—1-VIII-95 (95G4817). 

Aphis nr salicariae Koch [on Epilobium angustifolium]: WA, CHELAN—Swauk Pass, 1-VIII-95 (95G4927+); 
YAKIMA—YIR, Old Mead Canyon, 25-VII-95 (95G442). 

Aphis varians Patch [on Epilobium angustifolium]: ID, KOOTENAI—Rathdrum, 29-VII-95 (95G4437+); WA, 
CHELAN—Stevens Pass, 16-IX-95 (95K134+#); KittrraAs—Lost Lk, 24-VII-95 (95G399+¢#). 

Aphis nr varians Patch: WA, CHELAN—1-VIII-95 on Epilobium angustifolium (95G48 17). 

Aphis sp.: WA, CHELAN—Chelan SP, 11-IX-97 on Salix sp. (97K042+); GARFIELD—Bear Cr Rd, 14-IX-95 on 
Lilium columbianum (951577); Kirtiras—Taneum Lk, 11-[X-97 on Pedicularis racemosa (97G4117#); 
WHITMAN—Pullman, 12-VIII-96 on Helianthus annuus (96T438); YAKIMA—YIR, Windy Point, 19-[X-95 
on Cimicifuga laciniata (95G631). 

Brachycaudus helichrysi (Kaltenbach): WA, YAKIMA—Wenas, 11-VI-97 on Prunus domestica (97G079). 

Brachycaudus tragopogonis (Kaltenbach): WA, WHITMAN—Pullman, 12-IX-95 on Tragopogon dubius 
(95T561). 

Cavariella aegopodii (Scopoli): WA, YAkIMA—nr Wenas Lk, 17-VI-97 on Lomatium sp. (97GO95+) & 
Umbelliferae (97G097). 

Cavariella pastinacae (L.): WA, YAKIMA—N. Fork Tieton River, 27-VI-96 on Heracleum lanatum (96G288). 

Diuraphis noxia (Kurdjumov) [on Triticum aestivum]: WA, AsoTiIN—Asotin Cr Rd, 20-VI-95 (95T161); 
BENTON [*]; KLIckIrat—Bickleton, 6-VIII-97 (97G354); SPOKANE—Mt Spokane SP, 22-VII-95 (95T495). 

Dysaphis plantaginea (Passerini) [on Pyrus malus]: WA, CHELAN—-WSU-Wenatchee, 30-V-96, 13-VI-96 
(96RO009, 96R028); YAKIMA—Grandview, 26,28-VI-96 (96G282, 96G308). 

Ericaphis gentneri (Mason): WA, BENTON—WSU-Prosser, 24-VI-97 on Crataegus sp. (97G124). 

Hyperomyzus lactucae (L.): WA, WHITMAN—Pullman, 12-IX-95 on Lactuca serriola (95T559). 

Hyperomyzus nigricornis (Knowlton): WA, GARFIELD—Alder Thicket, FR 40, 22-VIII-95 on Senecio sp. 
(95T447); KirrirAs—Ellensburg, 13-VI-97 on Ribes sp. (97G084). 

Macrosiphum euphorbiae (Thomas): ID, KOOTENAI—Rathdrum, 29-VII-95 on Epilobium angustifolium 
(95G4437+); WA, KittirAs—Lost Lk, 6-VIII-97 on Castilleja (97G342+); YAKIMA [on Solanum 
lycopersicon* }. 

Macrosiphum gaurae (Williams): WA, YAKIMA—Wenas Lk, 11-VI-97 on Madia sp. (97G059). 

Macrosiphum pallidum (Oestlund): WA, YAKIMA—American River, 5-IX-97 on Epilobium minutum 
(97G3757). 

Macrosiphum valerianae (Clarke): WA, Kittiras—Lost Lk, 24-VII-95 on Epilobium angustifolium 
(95G399F +). 

Macrosiphum sp. [nr fuscicornis MacDougall & pallidum (Oestlund)]: WA, CHELAN—Swauk Pass, 1-VIII-95 
on Epilobium angustifolium (95G4927). 

Metopolophium dirhodum (Walker): WA, YAKIMA—Glade Rd, 7-VII-97 on Triticum aestivum (97G2177). 

Myzodium knowltoni Smith & Robinson: WA, KLickirat—John Day Dam, 28-IV-95 on Veronica anagallis- 
aquatica (95GO0097F +). 

Myzus cerasi (FE): WA, CHELAN—Wenatchee Heights, 6-VI-96 on Prunus sp. (96RO19). 

Myzus persicae (Sulzer): WA, DoUGLAs—Orondo, 25-V-95 on Prunus armeniaca (95RO11); KLickIrat—John 
Day Dam, 28-IV-95 on Veronica anagallis-aquatica (95G009+); WHITMAN—Pullman, 13-VI-96 on 
unknown plant (96T0317). 

Myzus sp.: WA, WHITMAN [on Forsythia sp.*]. 

Nasonovia ribisnigri (Mosley): WA, PIERCE—RNP, Tipsoo Lk, 5-[X-97 on Castilleja sp. (97G3617). 

Nasonovia (Kakimia) sp.: WA, GARFIELD—Spruce Spring, 22-VII-95 on Castilleja sp. (95T4637+); 
KitriraAs—Lost Lk, 6-VIII-97 on Castilleja sp. (97G342+); WHITMAN—Kamiak Butte, 12-IX-95 on 
Penstemon sp. (95T551). 

Nearctaphis bakeri (Cowen): WA, AsoTIN—De Spain Springs, 1-VIII-95 on Penstemon sp. (95T383). 

Nearctaphis kachena (Hottes) [on Castilleja sp. unless otherwise indicated]: ID, BENEWAH—McCroskey SP, 
31-VIII-95 (951522); WA, GaRFIELD—Spruce Spring, 22-VII-95 (95T4637£); Teal Springs, 22-VIII-96 
(96T484); Kirtiras—Lost Lk, 6-VIII-97 (97G3427+); Taneum Lk, 11-IX-97 on Pedicularis racemosa 
(97G411+£); SPOKANE—Mt Spokane SP, 24-VIII-95 (95T479, 95T485); YAKIMA—YIR, Mt Adams Lk Rd, 
23-VI-95 (95G242). 

Phorodon humuli (Schrank) [on Prunus salacina unless otherwise indicated]: WA, BENTON—-WSU-Prosser, 


728 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


19,25-IV-95, 1-V-95 (95G006, 95G007, 95GO016), 9-IX-95 on Humulus lupulus (95K154); YAKIMA— 
Sunnyside, 24-V-96 on H. lupulus (96G079+); Yakima Arboretum, 30-V-96 (96G104). 

Rhopalosiphum insertum (Walker): WA, CHELAN—-WSU-Wenatchee, 30-V-96 on Pyrus communis (96RO15), 
17-V-95 on Pyrus malus (95RO07); DouGLAs—Orondo, 25-V-95 on Pyrus communis (95RO10), 11 & 28- 
V-96 on Pyrus malus (96RO03, 96RO004, 96X005); Bridgeport, 12 & 17-VI-96 on Pyrus malus (96X006, 
96X0077+); KittirAas—Ellensburg, 17-V-96 on Crataegus columbiana (96G066). 

Rhopalosiphum maidis (Fitch): WA, BENTON [on Panicum milliaceum*]; YAKIMA—Sunnyside, 2-X-95 on 
Hordeum vulgare (95G6717). 

Rhopalosiphum padi (L.): WA, BENTON [on Triticum aestivum & Hordeum vulgare* |—WSU-Prosser, 24-IV- 
92, on T. aestivum (92A010); GRANT—Ephrata, 8-VII-97, on Zea mays (97K058); YAKIMA—Sunnyside, 
2-X-95 on H. vulgare (95G6717). 

Sitobion avenae (FE) [on Triticum aestivum unless otherwise indicated]: WA, AsoTIN—De Spain Springs, 24- 
VII-96 on Aquilegia formosa (961339); BENTON—West Richland, 28-VI-95 (95G279); WSU-Prosser, 3, 
11 & 18-VII-97 (97F191, 97F192, 97F215, 97F217, 97F227, 97F243, 97F249, 97F253, 97F256); 
YAKIMA—Glade Rd, 7-VII-97 (97G2177). 

Uroleucon sp.: ID, BENEWAH—McCroskey SP, 31-VHI-95 on unknown plant (95T520); KOOTENAI—Rathdrum, 
29-VII-95 on Epilobium angustifolium (95G4437). 

Undetermined spp.: WA, WHITMAN—Pullman, 19-VI-96 on Pyrus malus (96T042). 


Praon nr unicum Smith 
Euceraphis gillettei Davidson: WA, Kitriras—Manastash Rd, 25-VI-97 on Alnus sp. (97G1347#). 
Oestlundiella flava (Davidson): WA, KittrrAs—Manastash Rd, 25-VI-97 on Alnus sp. (97G1347#). 


Praon yakimanum Pike & Stary 

Cavariella pastinacae (L.): WA, ASoTIN—Asotin Cr, 11-VII-95 on Oenothera (95T196). 

Diuraphis noxia (Kurdjumoyv) [on Triticum aestivum]: WA, Asotin—Pataha Cr Rd, 10-VIII-95 (95T435); 
BENTON [*]|—WSU-Prosser, 23-V-97 (97F100, 97F112); KLICKIrAat—Bickleton, 9-VII-95 (95G3197). 

Metopolophium dirhodum (Walker): WA, YAKIMA—Sunnyside, 28-VI-95 on Phalaris arundinacea (95G254). 

Rhopalosiphum padi (L.): WA, BENTON [on Triticum aestivum & Hordeum vulgare*|; GRANT—Ephrata, 8- 
VII-97 on Zea mays (97K058). 

Sitobion avenae (F.): WA, KLICKIrATt—Bickleton, 9-VH-95 on Triticum aestivum (96G3197). 

Undetermined sp.: WA, SPOKANE—Turnbull National WLA, 13-VI-95 on Amelanchier alnifolia (95T414). 


Praon spp. 

Acyrthosiphon lactucae (Passerini) [on Lactuca serriola]: WA, Kitt1rAs—S Fork Manastash Cr, 24-VII-95 
(95G376); Ellensburg, 22-VI-96 (96G266); YAKIMA—Grandview, 20-X-95 (95G728). 

Acyrthosiphon pisum (Harris): MT, SANDER [on Lathyrus odoratus*]; WA, YAKIMA—YIR, Delaney Springs, 
25-VII-95 on Vicia sp. (95G408). 

Amphorophora rubi (Kaltenbach): WA, KLIick1IrAt—Trout Lk, 29-VI-97 on Rubus sp. (97G168). 

Amphorophora sensoriata Mason: WA, YAKIMA—Grandview, 22-VII-95 on Rubus sp. (95G368). 

Aphis armoraciae Cowen: WA, KiTTITASs [on Mertensia paniculata*}. 

Aphis ceanothi Clarke: WA, KitriraAs—S Fork Manastash Cr, 25-VI-97 on Ceanothus velutinus (97G137). 

Aphis coweni Palmer: WA, KiTTiITAs [on Veratrum viride*]. 

Aphis fabae Scopoli: WA, Kirriras—Lost Lk, 6-VIII-97 on Senecio sp. (97G335). 

Aphis pomi de Geer: WA, YAKIMA [on Pyrus malus*}]. 

Aphthargelia symphoricarpi (Thomas) [on Symphoricarpos sp.]: WA, Kittitas [*]; KLIckIrAat—Trout Lk, 29- 
VI-97 (97G159). 

Aspidophorodon longicauda (Richards): WA, SKAMANIA [on Spiraea sp. *]. 

Cavariella aegopodii (Scopoli): WA, GARFIELD—nr FR40 & FR44, 22-VIII-95 on Osmorhiza chilensis 
(95T464+4). 

Ceruraphis viburnicola (Gillette): WA, SKAMANIA [on Carex sp.*]. 

Chaetosiphon fragaefolii (Cockerell): ID, BOUNDARY [on Rosa sp.¢*]; WA, PEND OREILLE—Le Clerc WLA, 
3-VUI-95 on native Rosa sp. (95T398); WA, STEVENS—Little Pend Oreille National WLA, 9-VII-96 on 
native Rosa sp. (96T2097¢#). 

Epameibaphis atricornis G&P: WA, BENTON—-WSU-Prosser, 4-V-95 on Chrysothamnus nauseosus 
(95G0207 +). 

Ericaphis gentneri (Mason): WA, KitrirAs—Colockum Pass, 2-VII-97 on Crataegus sp. (97G214). 

Ericaphis scammelli (Mason): STEVENS—Little Pend Oreille National WLA, 9-VII-96 on native Rosa sp. 
(96T209F£). 

Hyperomyzus lactucae (L.): WA, BENTON [on Sonchus oleraceust*]. 


VOLUME 102, NUMBER 3 q29 


Hyperomyzus nigricornis (Knowlton): WA, GARFIELD—Wickiup CG, 29-VIII-95 on Hieracium sp. (95T508); 
KittirAs—Ellensburg, 23 VI—-96 on Ribes sp. (96G267). 

Hyperomyzus sp.: WA, BENTON [on Sonchus oleraceus*]. 

Illinoia nr crystleae (Smith & Knowlton): WA, Kitriras—S Fork Manastash Cr Rd, mi marker 8, 16-VIII- 
96 on Lonicera involucrata (96G489). 

Illinoia maxima (Mason): WA, KLickirAt—Trout Lk, 29-VI-97 on Rubus parviflorus (97G164). 

Illinoia rhododendri (Wilson): WA, KittiTAs [on Rhododendron albiflorum*}). 

Illinoia sp.: WA, GARFIELD—Stentz Springs, 1-VIII-95 on Thalictrum sp. (95T367); PIERCE—RNP, nr Tipsoo 
Lk, 16-VIII-95 on Spiraea densiflora (95A037); SPOKANE—Mt Spokane SP, 27-VII-95, on Vaccinium sp. 
(951346). 

Macrosiphoniella ludovicianae (Oestlund): WA, AsoTIN—Couse Cr Rd, mi marker 5, 18-VII-95 on Artemisia 
ludoviciana (951257). 

Macrosiphoniella tanacetaria (Kaltenbach): WA, SPOKANE—nr Mt Spokane, 29-VI-95 on Tanacetum vulgare 
(95T1727%). 

Macrosiphum albifrons Essig [on Lupinus sp.]: WA, KING—Stampede Pass, 8-VIII-96 (96G444); KLICKITAT— 
Trout Lk, 11-VII-95 (95K068). 

Macrosiphum californicum (Clarke): WA, SPOKANE—Mt Spokane SP, 27-VII-95, on Salix sp. (95T348). 

Macrosiphum euphorbiae (Thomas): ID, LAaTtAH—Moscow Mtn, 3-VIII-96 on Castilleja sp. (96T398); WA, 
GARFIELD—nr FR40 & FR44, 22-VIII-95 on Osmorhiza chilensis (95T464++); SPOKANE—nr Mt Spokane, 
29-VI-95 on Tanacetum vulgare (96T172+); YAKIMA [on Chenopodium album+*|—N Fork Tieton River, 
15-VI-96 on Hieracium albertinum (96K094). 

Macrosiphum nr euphorbiae (Thomas): WA, YAKIMA—YIR, Mill Cr Guard Station, 6-VI-95 on Lomatium 
nudicaule (95G115). 

Macrosiphum nr gaurae (Williams) [det. by A. Jensen]: WA, YAKIMA—Wenas Lk, 11-VI-97 on Mentzelia 
laevicaulis (97GO52). 

Macrosiphum pallidum (Oestlund): WA, SPOKANE—Cheney, 23-V-95 on native Rosa sp. (95KO006). 

Macrosiphum rosae (L.): WA, WHITMAN—Pullman, 1-VII-96 on cultivated Rosa sp. (961376); YAKIMA [on 
Rosa sp.*]. 

Macrosiphum sp.: WA, YAKIMA—YIR, Howard Lk, 19-[X-95 on Pteridium aquilinum (95G640). 

Macrosiphini [immatures]: WA, BENTON—WSU-Prosser, 27-X-95 on Malva neglecta (95K2127). 

Metopolophium dirhodum (Walker): WA, YAKIMA—Sunnyside, 28-VI-95 on Phalaris arundinacea (95G254). 

Microsiphoniella artemisiae (Gillette): WA, AsoTiIn—Asotin Cr HMU, 5-VII-96 on Artemisia ludoviciana 
(96T141). 

Myzus cerasi (EF) [on Prunus avium]: WA, BENTON [*] WHITMAN—Pullman, 5-VII-96 (96T173). 

Myzus persicae (Sulzer): WA, BENTON—-WSU-Prosser, 20V—97 on Prunus persica (97G036). 

Myzus sp.: WHITMAN [on Forsythia sp.*]. 

Nasonovia nr cynosbati (Oestlund): WA, YAKIMA—N Fork Tieton River, 15-VI-96 on native Ribes sp. 
(96K091). 

Nasonovia polemonii (G&P): WA, Kittiras—Buck Meadows, 9-VI-96, on Polemonium pulcherrimum 
(96G347). 

Nasonovia (Kakimia) sp.. WA, SPOKANE—nr Peone, 29-VI-95 on Phlox sp. (95T176); YAKIMA—YIR, nr Mt 
Adams Lk, 4-VIII-95 on Lonicera cilosa (95KO091). 

Nearctaphis sp.: WA, ASOTIN—De Spain Springs, 11-VH-96 on Castilleja sp. (961236). 

Pleotrichophorus sp... WA, BENTON—-WSU-Prosser, 4-V-95 on Chrysothamnus nauseosus (95G020*%); 
KLICKITAT—Alderdale & Mabton-Bickleton Rds, 28-IV-95 on Chrysothamnus sp. (95GO12). 

Rhopalomyzus lonicerae (Siebold): WA, SPOKANE—nr Peone, 13-VII-95 on unknown Gramineae (95T228). 

Rhopalosiphum maidis (Fitch): WA, BENTON [on Echinochloa crus-gallit*}. 

Rhopalosiphum nymphaeae (L.): WA, YAKIMA—Morgan Lk, 28-VI-95 on Nymphaea sp. (95G262). 

Rhopalosiphum padi (L.): WA, BENTON [on Echinochloa crus-galli+*]. 

Sitobion avenae (EF) [on Triticum aestivum]: WA, BENTON—-WSU-Prosser, 25-VII-97 (97F276); YAKIMA— 
Zillah, 28-VI-95 (95G270). 

Uroleucon nr erigeronense (Thomas): WA, Kittiras—Buck Meadows, 9-VII-96 on Hieracium albiflorum 
(96G340). 

Uroleucon ivae Robinson: WA, YAKIMA [on /va xanthifolia*]. 

Uroleucon russellae (HRL): WA, SKAMANIA [on Anaphalis margaritacea* |. 

Uroleucon sonchi (L.): WA, BENTON—-WSU-Prosser, 6-VII-95 on Sonchus oleraceus (95A025). 

Uroleucon sp.: WA, YAKIMA—Union Gap, 23-X-95 on Centaurea maculosa (95G720). 

Undetermined spp.: MT, SANDERS [on Senecio sp.*]; OR, UMATILLA—Harris Pk, 11-VI-96 on Preridium 


730 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


aquilinum (96G235); WA, Asotin—nr Anatone, 30-V-95 on Philadelphus lewisii (95T042); De Spain 
Springs, 24-VII-96 on Compositae (96T346); KittrrAs—Quartz Mtn, 25-VII-96 on Rubus lasiococcus 
(96G407): KLickirat—nr Cleveland, 7-VII-97 on Lotus pinnatus (97G233); WHITMAN—Pullman, 9-VII- 
96 on Lactuca serriola (961228); 25-VII-95, on Lapsana communis (951332); YAKIMA—N Fork Tieton 
River, 27-VI-96 on Achlys triphylla (96G289). 


Genus PSEUDOPRAON Stary 


Pseudopraon mindariphagum Stary 
Mindarus abietinus Koch: WA, YAKIMA—Yakima Arboretum, 30-V-96, on Abies concolor (96G102). 
Mindarus sp.: WA, YAKIMA—Grandview, 10-VI-96, on Abies grandis (96G215); N. Fork Tieton River, 15- 
VI-96, on Abies sp. (96K0987). 


Genus TOXARES Haliday 


Toxares deltiger (Haliday) 
Macrosiphum adianti (Oestlund): OR, BENTON—McDonald State Forest, on unknown plant, 21-V-91 (91JO02). 
Macrosiphum walkeri Robinson [det. by A. Jensen]: OR, Hoop RIveR—Starvation Cr, 21-IX-95 on 
Polypodium glycyrrhiza (95K180). 


Genus TRIOXYS Haliday 


Trioxys artemisiarum Pike & Stary 
Aphis (Zyxaphis) canae Williams: WA, DouGLas—Badger Mtn, 13-VII-95, on Artemisia tridentata 
(95B0127#). 
Macrosiphoniella ludovicianae (Oestlund) [on Artemisia ludoviciana]: WA, Asotin [*]—Asotin Cr WLA, 5- 
VII-96 (96T 138). 
Microsiphoniella artemisiae (Gillette): WA, AsoTIN—Asotin Cr HMU, 5-VII-96, on Artemisia ludoviciana 
(96T141). 


Trioxys bonnevillensis Smith 

Aphis canae Williams [on Artemisia tridentata]: WA, DouGLAs—Badger Mtn, 13-VII-95 (95B0127); 
FRANKLIN—Kahlotus Hwy, 11-VI-96 (96G2427¢#). 

Artemisaphis artemisicola (Williams) [on Artemisia tridentata]: WA, BENTON—Six Prong Rd, 24-I'V-97 
(97G008+); FRANKLIN—Kahlotus Hwy, 11-VI-96 (96G2427+); YAKIMA, Glade Rd, 18-VI-97 (97G1037¢). 

Epameibaphis atricornis G&P [on Artemisia tridentata]: WA, BENTON [*]—ALE, 19-V-95 (95G072+); Byron 
Rd, 14-V-97 (97G0247+); KLICKITAT [*]; YAKIMA [7*]. 

Epameibaphis utahensis Knowlton & Smith: WA, KLICKITAT [on Artemisia tridentatat*]. 

Epameibaphis sp.: WA, BENTON—ALE, 8-V1-95 on Artemisia tridentata (95G1517). 

Flabellomicrosiphum knowltoni Smith [on Artemisia tridentata}: WA, BENTON [+*]; YAKIMA [+*]—YIR, Fort 
Simcoe, 30-V-95 (95G091). 

Flabellomicrosiphum tridentatae (Wilson) [on Artemisia tridentata]: WA, FRANKLIN [*]—Juniper Sand Dunes, 
23-V-95 (95GO88+, 95G089+); KLickiTaT [+*]—Alderdale Cr & Columbia River Rds, 28-IV-95 (95G013). 

Flabellomicrosiphum sp. [on Artemisia tridentata]: WA, KLICKITAT [+*] & YAKIMA [**]. 

Microsiphoniella acophorum (Smith & Knowlton) [on Artemisia tridentata]: WA, FRANKLIN [*]—Juniper Sand 
Dunes, 23-V-95 (95G088+, 95G0897); FRANKLIN [{*]. 

Microsiphoniella sp. (Smith & Knowlton): WA, BENTON [on Artemisia tridentata }*}. 

Obtusicauda artemisiphila (Knowlton & Allen): WA, BENTON—ALE, 10-V-95 on Chrysothamnus nauseosus 
(95G033+); KLICKIrTAT—nr Roosevelt [on Artemisia tridentata+*]. 

Obtusicauda coweni (Hunter): WA, AsoTIN—Asotin Cr HMU, 7-V-96 on Artemisia tridentata (9611427). 

Obtusicauda filifoliae (G&P) [on Artemisia tridentata): WA, BENTON—Byron Rd, 14-V-97 (97G0247#); nr 
Prosser, Rotha Rd, 2-V-95 (95G0O17+#); KLICKITAT [F£*]; YAKIMA [+=*]. 

Obtusicauda sp. {on Artemisia tridentata]: WA, BENTON [+£*]; KLIcKITaT [t£*]; YAKIMA [++*]. 

Pleotrichophorus quadritrichus (Knowlton & Smith)?: WA, YAKIMA—Wenas Lk, 11-VI-97 on Artemisia 
tridentata (97GO58t¢£). 

Pleotrichophorus sp. [on Artemisia tridentata]:. WA, BENTON [t£*]; KLickrrat—Alderdale Rd, 9-V-95 
(95G0287#). 

Pseudoepameibaphis essigi Knowlton & Smith: WA, KLICKITAT [on Artemisia tridentata +*]. 

Pseudoepameibaphis tridentata (Wilson) [on Artemisia tridentata]: WA, BENTON [+*]—ALE, 19-V-95 
(95G072+); Byron Rd, 14-V-97 (97G024++); KLickiTaT [+*]—Alderdale Rd, 9-V-95 (95G0287); KitTITAs 


VOLUME 102, NUMBER 3 731 


[+*]; YAKIMA [+*]—Naught Rd, 18-VI-97 (97G103++, 97G120); Wenas Lk, 11-VI-97 (97G0587); YIR, 
Ft. Simcoe, 30-V-96 (95G0917). 
Trioxys cirsii (Curtis) 
Drepanosiphum platanoidis (Schrank): WA, BENTON [on Acer pseudoplatanus*]. 
Trioxys complanatus Quilis 
Therioaphis riehmi (Borner) [on Melilotus sp.]: WA, BENTON—-WSU-Prosser, 11 & 18-VII-97 on Melilotus 
officinalis (97F218+, 97F254+); KLickIrat—Roosevelt, 24-IV-96 (96G014+); YAKIMA, YIR, Signal Peak, 
19-IX-95 (95G6257). 
Therioaphis sp.: WA, BENTON—-WSU-Prosser, 28-VI-95 on Medicago sativa (95G2777+). 
Trioxys pallidus (Haliday) [group] 
Chromaphis juglandicola (Kaltenbach) [on Juglans regia]: WA, BENTON—-WSU-Prosser, 6-[X-95 (95K 1437); 
KLIcKITAaT—Bickleton, 27-VIII-96 (96G511); YAKIMA [*]—Yakima Arboretum, 26-IX-95 (95G657). 
Eucallipterus tiliae (L.) [on Tilia americana]: MT, LINCOLN—Libby, 29-VII-95 (95G449); WA, BENTON— 
Prosser, 1-VIII-95, 25-IX-95 (95A029, 95G647, 95G648); WHITMAN—Pullman, 9-X-96 (96T586). 
Monelliopsis caryae (Monell) [on Juglans nigra unless otherwise indicated]: WA, BENTON [on J. regia*]; 


WSU-Prosser, 6-I[X-95 (95G6507); YAKIMA [*]—Yakima Arboretum, 7-VII-94 & 26-IX-95 (94K020, 
95G656). 

Myzocallis coryli (Goeze): WA, CLARK—WSU-Vancouver, 21-I[X-95 on Corylus sp. (95K176). 

Myzocallis multisetis Boudreaux & Tissot: WA, BENTON—-WSU-Prosser, 9-[X-95 on Quercus borealis 
(95K158). 

Myzocallis sp.: WA, BENTON [on Quercus rubra*|; WHITMAN—Pullman, 9-X-96 on Quercus sp. (96T579). 

Panaphis juglandis (Goeze) [on Juglans regia}: WA, BENTON [*]—Prosser 25-IX-95 (95G646); WSU-Prosser, 
6-IX-95 (95K1437); YAKIMA [*]—Grandview, 25-IX-95 (95G653). 


Trioxys rosae Pike & Stary 
Macrosiphum pallidum (Oestlund): WA, SPOKANE—nr Cheney, 23-V-95 on native Rosa sp. (95KO006). 
Macrosiphum rosae (L.): WA, YAKIMA [on native Rosa sp.*]. 


Trioxys setaceus Pike and Stary, new species 
(Figs. 51-55) 

Diagnosis.—The new species is similar to T. artemisiarum Pike and Stary, from which 
it may be distinguished as follows: head prevalently dark brown; antenna 1 1-segmented; 
metasomal tergum 1 parallel-sided, not dilated to apex; prong dorsal surface with 6 
perpendicular setae and 2 shorter setae close to the base; last sternum with a group or 
longitudinal row of setae equal in length with 2 neighboring setae at dorsal base of prong. 

Etymology.—The name of the new species is derived from the presence of numerous 
setae on the prong of the last metasomal sternum. 

Description.—Female. Head: Eye large. Malar space equal to /% eye length. Tentorio- 
ocular line ¥, intertentorial line. Antenna 11-segmented, reaching to mid-metasoma. 
Flagellomeres F, to F, approximately equal in size, F, to F; as long as wide. Mesosoma: 
Mesosoma smooth, with a simple row of sparse individual setae tracing the effaced notauli 
on the disc. Propodeum (Fig. 52) smooth, with indications of short carinae or longitudinal 
rugosites in the distal portion. Forewing (Fig. 53): Stigma triangular, length 3 width; 
distal abscissa of R1 (metacarpus) short, distinctly shorter than stigma width. Metasoma: 
Metasomal tergum 1 (Fig. 51) length almost 2X width at spiracles (spiracles slightly 
prominent), roughly parallel sided from spiracle to posterior margin. Genitalia (Fig. 54): 
Prong straight, except for slight curve apically, apex with 2 longitudinally dilated setae 
(Fig. 55). Prong dorsal setae perpendicular (6 setae about 2 prong diameter, 2 near base 
short equal to prong diameter and to setae on distal part of last sternum); prong ventral 
setae numerous, oblique, about equal to prong diameter. Coloration: Head dark brown; 
lower part, clypeus, and gena yellow, palpi light yellow. Antenna scape, pedicel, and base 
of flagellomere 1 yellow, remaining antenna brown. Legs yellow brown, trochanters and 
base of tibia somewhat lighter. Hind leg with dark tinge. Metasomal tergum 1 yellow, 


132 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 51-55. Trioxys setaceus, 2 [illustrations not to equal scale]. 51, Metasomal tergum 1. 52, Propodeum. 
53, Fore wing, in part. 54, Genitalia. 55, Prong, close-up of apex. Abbreviations: R1, distal abscissa of post- 
marginal vein [= metacarpus]. 


terga 2 and 3 with yellow center and brown lateral spots; middle terga brown; distal 
tergum yellow. Ovipositor sheath yellow with brown apex; prong completely yellow to 
brownish at apex. Length of body: about 1.9—2.2 mm. 

Male.—Antenna 13-segmented. Coloration: head brown; clypeus and mandibles yellow, 
palpi light brown. Mesosoma brown. Wings hyaline, venation brown. Legs light brown, 
trochanter and base of tibia yellow. Metasoma brown, except tergum | yellow. 

Holotype.—?, USA, WA, AsoTIN Co., Schumaker Rd, 18-VII-1995, T. Miller collector 
(collection code 95T266). Host aphid: Acyrthosiphon macrosiphum (Wilson) on 
Amelanchier alnifolia. Deposited in USNM. 

Paratypes.—S specimens, collection data same as holotype, deposited in collection of 
WSU-Prosser (4 2, 1 3). 

Other material reared from: 

Acyrthosiphon macrosiphum (Wilson) [on Amelanchier alnifolia, unless indicated 
otherwise]: MT, LINCOLN—Libby, 6-VII-96 (96G322); WA, AsoTiIN—Anatone, 18-VII-95 
(95T266, 13 specimens); PEND OREILLE—Le Clerc WLA, 3-VIII-95 (95T402); SPOKANE, 
Cooper Rd nr Hwy 206, 29-VI-1995 (95T180, 5 specimens); nr Mt Spokane, on Ribes 
sp.?, (95T174). 


VOLUME 102, NUMBER 3 133) 


Trioxys tenuicaudus Stary 
Eucallipterus tiliae (L.): WA, BENTON—Prosser, 25-IX-95 on Tilia americana, (95G647, 95G648); 
KLICKITAT—Trout Lk, 29-VI-97 on T. sp. (97G174); WHITMAN—Pullman, 14-VIII-96 on T. americana, 
(96T443). 


Trioxys spp. 

Aphis coweni Palmer: WA, KitTiTAs [on Veratrum viride*]. 

Chaetosiphon fragaefolii (Cockerell): WA, SPOKANE—Turnbull National WLA, 1-VI-95, on native Rosa sp. 
(95T048). 

Eucallipterus tiliae (L.): WA, WHITMAN—Pullman, 1-VIII-96 on Tilia americana (961377). 

Euceraphis sp.: WA, SKAMANIA—Yellow Jacket Cr, 25-IX-94 on Trautvetteria caroliniensis (94G0217#). 

Macrosiphoniella ludovicianae (Oestlund): WA, YAKIMA—Naches, 28-VII-97 on Artemisia ludoviciana 
(97G277). 

Macrosiphum rosae (L.): WA, YAKIMA—nr Yakima, 3-VI-94 on native Rosa sp. (94/063). 

Monelliopsis nr caryae (Fitch): WA, BENTON—-WSU-Prosser, 24-VI-97 on Juglans nigra (97G126). 

Myzocallis nr granovskyi Boudreaux & Tissot: WA, BENTON—-WSU-Prosser, 24-VI-97 on Quercus rubra 
(97G125). 

Oestlundiella flava (Davidson): WA, SKAMANIA—Yellow Jacket Cr, 25-IX-94 on Trautvetteria caroliniensis 
(94G0217+#). 

Panaphis juglandis (Goeze): WA, BENTON—-WSU-Prosser, 24-VI-97 on Juglans regia (97G127). 

Pleotrichophorus sp.: WA, YAKIMA [on Haplopappus resinosus*]. 


Genus XENOSTIGMUS Smith 


Xenostigmus bifasciatus (Ashmead) 
Cinara ponderosae (Williams) [on Pinus ponderosa]: WA, AsoTiIn—De Spain Spring, 5-VII-96 (96T149); 
YAKIMA [*]. 
Cinara sp.: WA, AsoTiN—Cooper Canyon, 18-VII-96 on Pinus ponderosa (96T308). 
Undetermined sp.: WA, YAKIMA—Clear Lk, 15-VI-96 on Pinus ponderosa (96K087). 


734 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


APHID HOST—PARASITOID INDEX 


+—INDICATES APHID-PARASITOID ASSOCIATION NOT CERTAIN 


Acyrthosiphon kondoi 
Aphidius ervi 
Acyrthosiphon lactucae 
Aphidius avenaphis + 
Aphidius ervi 
Aphidius polygonaphis 
Aphidius sp. 
Diaeretiella rapae 
Ephedrus californicus 
Lysiphlebus testaceipes 
Praon occidentale 
Praon unicum 
Praon sp. 
Acyrthosiphon macrosiphum 
Aphidius sp. 
Lysiphlebus testaceipes 
Praon humulaphidis 
Praon pequodorum 
Trioxys setaceus 
Acyrthosiphon pisum 
Aphidius ervi 
Aphidius nr pisivorus 
Aphidius pulcher 
Aphidius smithi 
Aphidius sp. 
Diaeretiella rapae 
Lysiphlebus testaceipes 
Monoctonus sp. 
Praon occidentale 
Praon pequodorum 
Praon sp. 
Amphorophora geranii 
Aphidius sp. 
Praon occidentale 
Amphorophora parviflori 
Aphidius polygonaphis 
Amphorophora rubi 
Praon pequodorum 
Praon sp. 
Amphorophora nr rubitoxica 
Aphidius sp. 
Amphorophora sensoriata 
Praon sp. 
Amphorophora sp.? 
Aphidius polygonaphis 
Anoecia corni 
Lysiphlebus testaceipes 
Aphis armoraciae 
Adialytus fuscicornis 
Aphidius sp. 
Lysiphlebus testaceipes 
Praon sp. 


Aphis nr armoraciae 
Lysiphlebus testaceipes 

Aphis canae 
Binodoxys clydesmithi = 
Lysaphidus adelocarinus 
Lysiphlebus testaceipes + 
Lysiphlebus utahensis ~ 
Trioxys artemisiarum + 
Trioxys bonnevillensis + 

Aphis ceanothi 
Aphidius polygonaphis 
Binodoxys carolinensis 
Lysiphlebus testaceipes 
Praon sp. 

Aphis nr ceanothi 
Lysiphlebus testaceipes 

Aphis nr chrysothamni 
Lysiphlebus testaceipes 

Aphis coweni 
Ephedrus sp. 
Lysiphlebus testaceipes 
Praon americanum 
Praon unicum 
Praon sp. 

Trioxys sp. 

Aphis craccivora 
Aphidius ervi 
Aphidius sp. 
Diaeretiella rapae 
Lysiphlebus testaceipes 
Praon occidentale + 
Praon unicum 

Aphis nr epilobiaria 
Lysiphlebus testaceipes 

Aphis fabae 
Aphidius matricariae 
Aphidius sp. 

Binodoxys grafi 
Diaeretiella rapae 
Ephedrus californicus 
Lysiphlebus testaceipes 
Praon sp. 

Aphis nr fabae 
Diaeretiella rapae 
Lysiphlebus testaceipes 
Praon unicum 

Aphis filifoliae 
Lysiphlebus testaceipes 

Aphis frangulae 
Lysiphlebus testaceipes 

Aphis nr frangulae 
Lysiphlebus testaceipes 


Aphis gossyptt 
Lysiphlebus testaceipes 
Aphis hederae 
Lysiphlebus testaceipes 
Aphis helianthi 
Adialytus fuscicornis 
Aphidius matricariae 
Aphidius polygonaphis + 
Aphidius sp. 
Binodoxys carolinensis 
Binodoxys grafi 
Diaeretiella rapae 
Ephedrus californicus 
Ephedrus clavicornis 
Lysiphlebus testaceipes 
Lysiphlebus sp. 
Praon occidentale 
Praon unicum 
Aphis nr helianthi 
Lysiphlebus testaceipes 
Aphis hermistonii 
Lysiphlebus testaceipes 
Aphis holodisci 
Aphidius sp. 
Binodoxys graft 
Ephedrus californicus 
Lysiphlebus testaceipes 
Aphis lugentis 
Lysiphlebus testaceipes 
Praon occidentale 
Aphis lupini 
Lysiphlebus testaceipes 
Aphis neogillettei 
Lysiphlebus testaceipes 
Aphis nerit 
Aphidius colemani 
Lysiphlebus testaceipes 
Aphis nigratibialis 
Aphidius sp. 
Binodoxys carolinensis 
Lysiphlebus testaceipes 
Praon unicum 
Aphis oenotherae 
Lysiphlebus testaceipes 
Praon unicum 
Aphis nr oenotherae 
Aphidius polygonaphis 
Binodoxys carolinensis 
Ephedrus californicus 
Ephedrus clavicornis 
Lysiphlebus testaceipes 


VOLUME 102, NUMBER 3 


Monoctonus washingtonensis 


Praon unicum 
Aphis pomi 


Lysiphlebus testaceipes + 


Praon unicum 
Praon sp. 

Aphis rumicis 
Lysiphlebus testaceipes 
Praon unicum 

Aphis salicariae 
Binodoxys carolinensis 
Binodoxys grafi 
Binodoxys sp. 
Lysiphlebus testaceipes 
Praon americanum 
Praon unicum 

Aphis nr salicariae 
Binodoxys grafi 
Lysiphlebus testaceipes 
Praon unicum 

Aphis sambuci 
Lysiphlebus testaceipes 

Aphis spiraecola 
Lysiphlebus testaceipes 

Aphis nr triglochinis 
Binodoxys grafi 
Lysiphlebus testaceipes 

Aphis nr valerianae 
Lysiphlebus testaceipes 

Aphis varians 
Aphidius sp. 
Binodoxys carolinensis 
Binodoxys grafi 
Ephedrus californicus 
Ephedrus clavicornis 
Ephedrus sp. 
Lysiphlebus testaceipes 
Praon unicum 

Aphis nr varians 
Binodoxys carolinensis 
Binodoxys grafi 
Ephedrus californicus 
Lysiphlebus testaceipes 
Praon unicum 

Aphis (Zyxaphis) sp. 
Aphidius sp. 
Lysaphidus adelocarinus 
Lysaphidus rosaphidis + 
Lysiphlebus testaceipes 

Aphis sp. n. 

Binodoxys carolinensis 
Lysiphlebus testaceipes 

Aphis spp. 

Aphidius ervi 
Binodoxys carolinensis 
Lysiphlebus testaceipes 
Lysiphlebus sp. 

Praon unicum 


++ 


Aphthargelia symphoricarpi 


Aphidius matricariae 
Ephedrus californicus 
Ephedrus clavicornis 
Lysaphidus sp. 
Lysiphlebus testaceipes 
Praon coniforme 
Praon sp. 
Artemisaphis artemisicola 
Binodoxys clydesmithi 


Lysaphidus adelocarinus 


Lysiphlebus utahensis 
Praon artemisicola 
Trioxys bonnevillensis 


Aspidophorodon longicauda 


Aphidius ervi 
Aphidius sp. 
Praon sp. 


Aspidophorodon nr longicauda 


Ephedrus lacertosus + 
Boernerina variabilis 
Betuloxys alnicolus ¢ 
Lysiphlebus testaceipes 
Brachycaudus cardui 
Aphidius matricariae 
Aphidius polygonaphist 
Diaeretiella rapae 
Ephedrus persicae 
Lysiphlebus testaceipes 
Brachycaudus helichrysi 
Aphidius colemani 
Aphidius ervi 
Aphidius matricariae 
Aphidius sp. 
Diaeretiella rapae 
Ephedrus californicus = 
Ephedrus persicae 
Lysiphlebus testaceipes 


Monoctonus campbellianus 


Praon occidentale 
Praon unicum 


Brachycaudus tragopogonis 


Aphidius sp. 
Diaeretiella rapae 
Lysiphlebus testaceipes 
Lysiphlebus sp. 
Praon unicum 
Brachycorynella asparagi 
Diaeretiella rapae 
Lysiphlebus testaceipes 
Braggia eriogoni 
Ephedrus californicus 
Lysiphlebus testaceipes 
Braggia nr. eriogoni. 
Lysiphlebus testaceipes 
Braggia sp. n. 
Lysiphlebus testaceipes 
Braggia sp. 


Diaeretiella rapae 
Lysiphlebus testaceipes 
Brevicoryne brassicae 
Aphidius ervi 
Aphidius matricariae 
Diaeretiella rapae 
Ephedrus californicus 
Lysiphlebus testaceipes 
Capitophorus elaeagni 
Aphidius matricariae 
Aphidius polygonaphis 
Aphidius sp. 
Ephedrus californicus 
Lysiphlebus testaceipes 
Praon occidentale + 
Carolinaia howardii 
Aphidius sp. 
Cavariella aegopodii 
Aphidius salicis 
Ephedrus californicus 
Lysiphlebus testaceipes 
Praon unicum 
Praon sp. £ 
Cavariella nr aegopodii 


Binodoxys carolinensis + 


Cavariella pastinacae 
Aphidius salicis 
Ephedrus californicus 
Lysiphlebus testaceipes 
Praon unicum 
Praon yakimanum 

Cedoaphis incognita 
Lysiphlebus flavidus 

Ceruraphis viburnicola 
Lysiphlebus testaceipes 
Praon sp. 

Chaetosiphon fragaefolii 
Aphidius sp. 

Binodoxys sp. 
Ephedrus californicus 
Ephedrus sp. ¢ 
Lysaphidus rosaphidis 
Lysaphidus sp. = 
Lysiphlebus testaceipes 
Praon sp. 

Trioxys sp. 

Chaetosiphon nr hottesi 
Aphidius ervi 
Aphidius sp. ¢ 

Chaetosiphon tetrarhodum 
Aphidius sp. = 

Chaetosiphon thomasi 
Praon occidentale 

Chaetosiphon nr thomasi 
Lysaphidus rosaphidis 

Chaitophorus populicola 
Adialytus salicaphis 
Lysiphlebus testaceipes 


785 


736 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Chaitophorus populifolii 
Adialytus salicaphis 
Ephedrus chaitophori 

Chaitophorus utahensis 
Adialytus salicaphis 

Chaitophorus viminalis 
Adialytus salicaphis 
Lysiphlebus testaceipes 

Chaitophorus sp. 
Adialytus salicaphis 

Chromaphis juglandicola 
Trioxys pallidus 

Cinara brevispinosa 
Pauesia sp. 

Cinara chinookiana 
Lysiphlebus testaceipes 

Cinara pilicornis 
Pauesia juniperaphidis 

Cinara ponderosae 
Pauesia ahtanumensis 
Pauesia pahtonis 
Pauesia ponderosaecola 
Pauesia sp. 
Xenostigmus bifasciatus 

Cinara pseudotaxifoliae 
Pauesia pseudotsugae 

Cinara pseudotsugae 
Pauesia sp. 

Cinara spp. 

Lysiphlebus testaceipes 
Pauesia ahtanumensis 
Pauesia bicolor 
Pauesia sp. 
Xenostigmus bifasciatus 

Diuraphis noxia 
Aphidius avenaphis 
Aphidius ervi 
Aphidius matricariae 
Aphidius sp. 
Diaeretiella rapae 
Ephedrus californicus 
Ephedrus sp. 
Lysiphlebus testaceipes 
Monoctonus washingtonensis 
Praon gallicum 
Praon occidentale 
Praon unicum 
Praon yakimanum 

Drepanosiphum platanoidis 
Trioxys cirsii 

Dysaphis plantaginea 
Aphidius matricariae + 
Diaeretiella rapae 
Ephedrus clavicornis 
Lysiphlebus testaceipes 
Praon occidentale 
Praon unicum 


Epameibaphis atricornis 
Lysaphidus adelocarinus 
Lysiphlebus utahensis + 
Praon sp. + 
Praon artemisicola 
Trioxys bonnevillensis 

Epameibaphis utahensis 
Trioxys bonnevillensis 

Epameibaphis sp. 

Trioxys bonnevillensis 

Ericaphis gentneri 
Aphidius nr hortensis 
Aphidius sp. 

Ephedrus lacertosus 
Lysaphidus sp. 
Praon unicum 
Praon sp. 

Ericaphis nr gentneri 
Aphidius sp. 

Lysiphlebus testaceipes 

Ericaphis scammelli 
Aphidius sp. 

Ephedrus sp. + 
Lysaphidus sp. ¢ 
Praon sp. = 

Ericaphis wakibae 
Aphidius polygonaphis 
Aphidius sp. 

Ephedrus californicus + 
Praon occidentale + 

Eriosoma americanum 
Lysiphlebus testaceipes = 

Eriosoma lanigerum 
Lysiphlebus testaceipes + 

Eucallipterus tiliae 
Trioxys pallidus 
Trioxys tenuicaudus 
Trioxys sp. 

Euceraphis gillettei 

Aphidius sp. $ 
Praon nr unicum ¢ 

Euceraphis sp. 

Trioxys sp. £ 

Flabellomicrosiphum knowltoni 
Lysaphidus adelocarinus 
Trioxys bonnevillensis 

Flabellomicrosiphum tridentata 
Lysaphidus adelocarinus + 
Lysiphlebus utahensis + 
Trioxys bonnevillensis 

Flabellomicrosiphum sp. 
Lysaphidus adelocarinus 
Praon artemisicola 
Trioxys bonnevillensis 

Forda marginata 
Adialytus fuscicornis 

Hayhurstia atriplicis 


Aphidius ervi 
Diaeretiella rapae 

Hyadaphis foeniculi 
Aphidius colemani 
Lysiphlebus testaceipes + 

Hyalopteroides humilis 
Aphidius ervi 

Hyalopterus pruni 
Aphidius ervi + 
Aphidius matricariae + 
Monoctonus campbellianus 

Hyperomyzus lactucae 
Aphidius ervi 
Aphidius segmentatus 
Ephedrus californicus 
Praon unicum 
Praon sp. 

Hyperomyzus nigricornis 
Aphidius ervi 
Aphidius polygonaphis 
Aphidius segmentatus 
Aphidius sp. 

Ephedrus californicus 
Ephedrus sp. 
Lysiphlebus testaceipes 
Praon occidentale 
Praon unicum 

Praon sp. 

Hyperomyzus spp. 
Aphidius polygonaphis 
Aphidius segmentatus 
Aphidius sp. 

Ephedrus californicus 
Praon occidentale 
Praon sp. 

Illinoia nr azaleae 
Aphidius polygonaphis 

Illinoia corylina 
Aphidius matricariae 
Diaeretiella rapae 

Illinoia crystleae bartholomewi 
Aphidius sp. 

Illinoia nr crystleae 
Aphidius sp. 

Praon sp. 

Illinoia davidsoni 
Aphidius ervi 
Aphidius polygonaphis 
Aphidius sp. 

Praon occidentale 

Illinoia maxima 
Ephedrus clavicornis 
Praon sp. 

Illinoia rhododendri 
Aphidius sp. 
Binodoxys sp. 
Monoctonus sp. 

Praon sp. 


VOLUME 102, NUMBER 3 


Illinoia richardsi 
Praon occidentale + 

Illinoia spiraeae 
Lysiphlebus testaceipes 

Ilinoia nx thalictri 
Praon occidentale 

Illinoia spp. 

Aphidius ervi 
Aphidius polygonaphis 
Aphidius sp. 

Ephedrus californicus 
Monoctonus sp. 

Praon humulaphidis 
Praon occidentale 
Praon nr occidentale 
Praon pequodorum 
Praon sp. 

Illinoia sp. ? 

Praon humulaphidis 

Liosomaphis berberidis 
Aphidius nr hortensis 
Aphidius sp. 

Lipaphis erysimi 
Aphidius avenaphis 
Aphidius colemani 
Diaeretiella rapae 

Macrosiphoniella ludovicianae 
Aphidius ohioensis 
Aphidius polygonaphis 
Aphidius sp. 

Ephedrus californicus 
Praon occidentale 
Praon sp. 
Trioxys artemisiarum 
Trioxys sp. 
Macrosiphoniella nr ludovicianae 
Praon occidentale 

Macrosiphoniella tanacetaria 
Aphidius ohioensis 
Ephedrus californicus 
Praon sp. = 

Macrosiphum adianti 
Toxares deltiger 

Macrosiphum albifrons 
Aphidius ervi 
Aphidius lupini 
Ephedrus californicus 
Ephedrus lacertosus 
Praon occidentale 
Praon sp. 

Macrosiphum californicum 
Aphidius sp. 

Ephedrus californicus 
Praon sp. 

Macrosiphum clydesmithi 
Aphidius nigripes 
Aphidius polygonaphis 
Aphidius sp. 


Macrosiphum creelii 
Aphidius ohioensis 
Ephedrus californicus 

Macrosiphum euphorbiae 
Aphidius colemani 
Aphidius ervi 
Aphidius matricariae 
Aphidius nigripes 
Aphidius polygonaphis 
Aphidius sp. 
Diaeretiella rapae 
Ephedrus californicus 
Ephedrus lacertosus 
Lysaphidus sp. 
Lysiphlebus testaceipes 
Monoctonus sp. 

Praon fulvum 
Praon humulaphidis 
Praon occidentale 
Praon unicum 
Praon sp. 

Macrosiphum nr euphorbiae 
Aphidius polygonaphis 
Diaeretiella rapae 
Ephedrus californicus 
Ephedrus sp. 

Praon sp. 

Macrosiphum gaurae 
Aphidius ervi 
Ephedrus californicus 
Lysiphlebus testaceipes 
Praon unicum 

Macrosiphum nr gaurae 
Praon sp. 

Macrosiphum osmaroniae 
Praon occidentale 

Macrosiphum pallidum 
Aphidius nr rosae 

Aphidius sp. = 
Ephedrus californicus + 
Praon unicum 

Praon sp. 

Trioxys rosae 

Macrosiphum nr pallidum 
Aphidius ervi + 
Lysiphlebus testaceipes + 

Macrosiphum pteridis 

Aphidius polygonaphis 

Macrosiphum rhamni 

Aphidius sp. 

Ephedrus sp. 

Praon simulans 
Macrosiphum rosae 

Aphidius ervi 

Aphidius polygonaphis 

Aphidius sp. 

Ephedrus californicus 

Praon occidentale 


UY 


Praon sp. 
Trioxys rosae 
Trioxys sp. 
Macrosiphum tuberculaceps 
Aphidius sp. 
Monoctonus pacificus 
Praon fulvum 
Macrosiphum valerianae 
Aphidius ohioensis 
Aphidius polygonaphis 
Ephedrus californicus + 
Praon unicum ¢ 
Macrosiphum walkeri 
Aphidius nr polygonaphis 
Monoctonus sp. 
Toxares deltiger 
Macrosiphum spp. 
Aphidius ervi 
Aphidius polygonaphis 
Aphidius sp. 
Diaeretiella rapae 
Ephedrus lacertosus 
Lysiphlebus testaceipes 
Praon fulvum 
Praon humulaphidis 
Praon occidentale 
Praon simulans 
Praon unicum 
Praon sp. 
Toxares deltiger 
Macrosiphum sp. ? 
Aphidius ohioensis 
Praon occidetale 
Macrosiphini [immatures] 
Ephedrus lacertosus 
Monoctonus pacificus 
Praon sp. 
Metopolophium dirhodum 
Aphidius ervi 
Aphidius rhopalosiphi 
Aphidius sp. 
Ephedrus californicus = 
Monoctonus sp. 
Praon gallicum 
Praon occidentale = 
Praon unicum 
Praon yakimanum 
Praon sp. 
Metopolophium sp. 
Ephedrus lacertosus ¢ 
Metopolophium sp. ? 
Praon occidentale 
Microlophium nr sibiricum 
Praon occidentale 
Microsiphoniella acophorum 
Trioxys bonnevillensis 
Microsiphoniella artemisiae 
Aphidius ohioensis 


738 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Lysaphidus adelocarinus 
Praon sp. 
Trioxys artemisiarum 
Microsiphoniella sp. 
Trioxys bonnevillensis 
Mindarus abietinus 
Pseudopraon mindariphagum 
Mindarus sp. 
Pseudopraon mindariphagum 
Monelliopsis caryae 
Trioxys pallidus 
Monelliopsis nr caryae 
Trioxys sp. 
Myzocallis coryli 
Trioxys pallidus 
Myzocallis nr granovskyi 
Trioxys sp. 
Myzocallis multisetis 
Trioxys pallidus 
Myzocallis sp. 
Trioxys pallidus 
Myzodium knowltoni 
Praon unicum <= 
Myzus ascalonicus 
Lysiphlebus testaceipes 
Myzus cerasi 
Aphidius sp. 
Ephedrus californicus 
Lysiphlebus testaceipes 
Praon unicum 
Praon sp. 
Myzus nr cerasi 
Lysiphlebus testaceipes 
Myzus nr ornatus 
Aphidius ervi ¢ 
Myzus persicae 
Aphidius colemani 
Aphidius ervi 
Aphidius matricariae 
Diaeretiella rapae 
Ephedrus californicus 
Lysiphlebus testaceipes 
Praon occidentale 
Praon unicum 
Praon sp. 
Myzus spp. 
Aphidius colemani 
Aphidius ervi 
Aphidius matricariae 
Diaeretiella rapae 
Praon unicum 
Praon sp. 
Nasonovia alpina 
Aphidius kakimiaphidis 
Nasonovia aquilegiae 
Aphidius kakimiaphidis 
Aphidius sp. 
Praon occidentale 


Nasonovia nr aquilegiae 
Aphidius sp. 

Nasonovia crenicorna 
Aphidius polygonaphis 

Nasonovia cynosbati 
Aphidius kakimiaphidis 

Nasonovia nr cynosbati 
Aphidius kakimiaphidis 
Praon sp. 

Nasonovia houghtonensis 
Aphidius polygonaphis 
Harkeria rufa 

Nasonovia nr houghtonensis 
Aphidius kakimiaphidis 

Nasonovia polemonii 
Aphidius sp. 

Praon occidentale 
Praon sp. 

Nasonovia ribisnigeri 
Ephedrus californicus 
Praon humulaphidis 
Praon unicum 

Nasonovia wahinkae 
Aphidius kakimiaphidis 

Nasonovia spp. 

Aphidius kakimiaphidis 
Aphidius sp. 

Ephedrus californicus 
Lysaphidus sp. 
Lysiphlebus testaceipes 
Praon occidentale + 
Praon unicum 

Praon sp. 

Nearctaphis bakeri 
Lysiphlebus testaceipes 
Praon occidentale + 
Praon unicum 

Nearctaphis californica 
Lysiphlebus testaceipes 

Nearctaphis kachena 
Lysiphlebus testaceipes + 
Praon occidentale 
Praon unicum 

Nearctaphis yohoensis 
Lysiphlebus testaceipes 

Nearctaphis spp. 
Lysiphlebus testaceipes 
Praon sp. 

Obtusicauda artemisiphila 
Ephedrus californicus 
Lysiphlebus utahensis + 
Trioxys bonnevillensis 

Obtusicauda coweni 
Binodoxys clydesmithi 
Binodoxys coruscanigrans 
Ephedrus californicus 
Lysaphidus adelocarinus 
Lysiphlebus utahensis 


Praon artemisaphis 

Trioxys bonnevillensis 
Obtusicauda filifoliae 

Lysiphlebus utahensis * 

Trioxys bonnevillensis = 
Obtusicauda spp. 

Ephedrus californicus 

Lysaphidus adelocarinus 

Lysiphlebus utahensis 

Praon artemisaphis 

Trioxys bonnevillensis + 
Oestlundiella flava 

Aphidius sp. += 

Praon nr unicum + 

Trioxys sp. 
Ovatus crataegarius 

Aphidius ervi 

Aphidius matricariae 

Aphidius sp. 

Ephedrus incompletus 
Panaphis juglandis 

Trioxys pallidus 

Trioxys sp. 
Periphyllus nr brevispinosus 

Adialytus salicaphis 
Periphyllus californiensis 

Euaphidius setiger 
Periphyllus lyropictus 

Euaphidius setiger 
Periphyllus sp. 

Adialytus salicaphis 

Euaphidius setiger 
Phorodon humuli 

Aphidius ervi ¥ 

Aphidius matricariae 

Binodoxys conei 

Diaeretiella rapae 

Lysiphlebus testaceipes 

Monoctonus campbellianus 

Praon occidentale 

Praon unicum 
Pleotrichophorus nr amsinckii 

Ephedrus californicus 

Lysaphidus adelocarinus 
Pleotrichophorus elongatus 

Lysaphidus ramithyrus 
Pleotrichophorus palmerae 

Lysaphidus adelocarinus 
Pleotrichophorus pseudoglandu- 

losus 

Ephedrus californicus + 
Pleotrichophorus quadritichus 

Praon artemiscola 

Trioxys bonnevillensis * 
Pleotrichophorus nr. wasatchi 

Lysaphidus ramithyrus 
Pleotrichophorus spp. 

Lysaphidus adelocarinus 


VOLUME 102, NUMBER 3 


Lysaphidus rosaphidis + 
Praon artemisicola + 
Praon sp. 
Trioxys bonnevillensis + 
Trioxys sp. 
Prociphilus sp. 
Aphidius sp. 
Ephedrus clavicornis 
Ephedrus sp. 
Pseudoepameibaphis essigi 
Lysaphidus adelocarinus 
Trioxys bonnevillensis 
Pseudoepameibaphis glauca 
Lysaphidus adelocarinus 
Pseudoepameibaphis tridentatae 
Lysaphidus adelocarinus 
Lysiphlebus testaceipes + 
Lysiphlebus utahensis + 
Praon artemisicola 
Trioxys bonnevillensis 
Pterocomma beulahense 
Euaphidius cingulatus 
Pterocomma bicolor 
Euaphidius cingulatus 
Pterocomma sanguiceps 
Euaphidius cingulatus 
Pterocomma smithiae 
Euaphidius cingulatus 
Pterocomma sp. 
Euaphidius cingulatus 
Lysiphlebus testaceipes 
Rhopalomyzus lonicerae 
Praon sp. 
Rhopalosiphum cerasifoliae 
Lysiphlebus testaceipes 
Rhopalosiphum enigmae 
Lysiphlebus testaceipes 
Rhopalosiphum insertum 
Aphidius matricariae 
Diaeretiella rapae 
Ephedrus californicus 
Lysiphlebus testaceipes 
Praon occidentale 
Praon unicum 
Rhopalosiphum maidis 
Aphidius ervi 
Diaeretiella rapae 
Ephedrus californicus + 
Lysiphlebus testaceipes 
Praon unicum 
Praon sp. 
Rhopalosiphum nymphaeae 
Diaeretellus palustris 
Lysiphlebus testaceipes 
Praon sp. 
Rhopalosiphum padi 
Aphidius avenaphis 
Aphidius ervi 


Aphidius matricariae 
Aphidius sp. 
Diaeretiella rapae 
Ephedrus californicus + 
Lysiphlebus testaceipes 
Monoctonus washingtonensis 
Praon gallicum + 
Praon occidentale 
Praon unicum 
Praon yakimanum 
Praon sp. 
Schizaphis graminum 
Lysiphlebus testaceipes 
Sipha elegans 
Adialytus ambiguus 
Sipha sp. 
Adialytus ambiguus 
Sitobion avenae 
Aphidius avenaphis 
Aphidius ervi 
Aphidius matricariae 
Aphidius rhopalosiphi 
Aphidius sp. 
Diaeretiella rapae 
Ephedrus californicus 
Ephedrus clavicornis 
Lysiphlebus testaceipes 
Praon gallicum 
Praon occidentale 
Praon unicum 
Praon yakimanum 
Praon sp. 
Sitobion fragariae 
Aphidius sp. 
Sitobion spp. 
Aphidius polygonaphis 
Ephedrus lacertosus 
Lysiphlebus testaceipes 
Therioaphis riehmi 
Praon exsoletum 
Praon pequodorum + 
Trioxys complanatus 
Therioaphis trifolii 
Praon exsoletum 
Therioaphis sp. 
Praon exsoletum 
Trioxys complanatus 
Tuberculatus columbiae 
Lysiphlebus testaceipes 
Uroleucon adenocaulonae 
Aphidius ohioensis 
Uroleucon nr arnesense 
Aphidius ohioensis 
Aphidius polygonaphis 
Ephedrus californicus 
Uroleucon cirsii 
Lysiphlebus testaceipes 


139 


Uroleucon nr epilobii 
Aphidius ohioensis 
Uroleucon erigeronense 
Aphidius ohioensis 
Aphidius segmentatus + 

Aphidius sp. 
Ephedrus californicus 
Praon occidentale 
Praon nr occidentale 

Uroleucon nr erigeronense 
Aphidius ohioensis 
Aphidius polygonaphis 
Ephedrus californicus 
Praon sp. 

Uroleucon escalantii 
Aphidius ohioensis 
Uroleucon nr escalantii 
Aphidius ohioensis 
Uroleucon gigantiphagum 
Aphidius ohioensis 

Uroleucon ivae 
Aphidius ohioensis 
Aphidius polygonaphis 
Aphidius sp. 
Diaeretiella rapae 
Ephedrus californicus 
Praon pequodorum 
Praon sp. 

Uroleucon nr ivae 
Praon nr occidentale 

Uroleucon katonkae 
Aphidius ohioensis 
Ephedrus californicus 

Uroleucon oliveri 
Aphidius ohioensis 

Uroleucon russellae 
Aphidius ohioensis 
Aphidius sp. 
Binodoxys rhagii 
Ephedrus californicus 
Praon sp. 

Uroleucon sonchi 
Aphidius ohioensis 
Aphidius sp. 

Ephedrus californicus 
Praon occidentale 
Praon pequodorum 
Praon sp. 

Uroleucon suzannae 
Aphidius ohioensis + 
Praon nr occidentale + 

Uroleucon zymozionense 
Praon occidentale 

Uroleucon spp. 
Acanthocaudus caudacanthus 
Aphidius ervi 
Aphidius ohioensis 
Aphidius sp. 


740 


Diaeretiella rapae 
Ephedrus californicus 
Lysiphlebus testaceipes 


Praon sp. 


Praon occidentale Aphidius sp. 


ACKNOWLEDGMENTS 


We express thanks to Washington State 
University collaborators W. Turner, G. 
Clevenger, J. Rumph, and T. Murray, and 
Wenatchee Valley College collaborator R. 
Gillespie for assistance in collecting aphids 
in the field and rearing parasitoids. We ex- 
press thanks to G. Ahlstrand and R. Lech- 
leitner for facilitating collections at Mt. 
Rainier National Park, and the Yakama In- 
dian Reservation for allowing access to 
their lands. This work was made possible 
with the support of numerous growers on 
whose land we frequently collected, and in 
part by funds provided by Washington State 
University, the Washington Wheat and Bar- 
ley Commissions, and the U.S. Department 
of Agriculture. 


LITERATURE CITED 


Gahan, A. B. 1911. Aphidiinae of North America. 
Maryland Agricultural Experiment Station Bulle- 
tin 152: 147-200. 

Huber, J. T. and M. J. Sharkey. 1993. Structure, pp. 
13-59. In Goulet, H. & J. T. Huber, eds., Hyme- 
noptera of the World: An Identification Guide to 
Families. Research Branch, Agriculture Canada, 
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. 

Liu, Ching-Shen. 1977. Genus Aphidius (Aphididae, 
Hym.) of California. Taiwan China, Special Bul- 
letin No. 11, 118 pp. 

Mackauer, M. and P. Stary. 1967. Ichneumonoidea, 
World Aphidiidae, pp. 1-195. Jn Delucchi, V and 
G. Remaudiere, eds., Index of Entomophagous In- 
sects. Le Francois, Paris. 

Marsh, P. M. 1979. Aphidiidae, pp. 295-313. Jn Krom- 
bein, K. V., BP D. Hurd, Jr., D. R. Smith, and B. 
D. Burks, eds., Catalog of Hymenoptera of Amer- 
ica North of Mexico, Vol. 1, Symphyta and Apo- 
crita (Parasitica). Smithsonian Institution Press, 
Washington, D.C. 


Praon unicum 


Utamphorophora humboldti 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Wahlgreniella nervata 
Aphidius polygonaphis 
Ephedrus californicus + 
Praon occidentale + 


Pike, K. S. and P. Stary. 1995. New species of parasitic 
wasps attacking cereal aphids in the Pacific North- 
west (Hymenoptera, Braconidae, Aphidiinae). 
Journal of the Kansas Entomological Society 68: 
408-414. 

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 (Hym.: Bra- 
conidae: Aphidiinae) from the Pacific Northwest, 
U.S.A. Proceedings of the Entomological Society 
of Washington 98: 570-591. 

Pike, K. S., P. Stary, G. Graf, D. Allison, L. Boydston, 
R. Miller, and T. Miller. 1997. Parasitoids (Hym., 
Braconidae, Aphidiinae) of aphids on big sage- 
brush (Artemisia tridentata) and prairie sage (Ar- 
temisia ludoviciana) in Washington State. Pro- 
ceedings of the Entomological Society of Wash- 
ington 99: 143-155. 

Pike, K. S., P. Stary, T. Miller, R. Miller, D. Allison, 
G. Graf, L. Boydston, and R. Gillespie. 1999. 
Ephedrus clavicornis sp. n. (Hymenoptera, Bra- 
conidae, Aphidiinae), an aphid parasitoid of the 
Pacific Northwest, and key to Nearctic Ephedrus. 
Journal of the Kansas Entomological Society 72: 
10-16. 

Remaudiére, G. and M. Remaudiére 1997. Catalogue 
of the World’s Aphididae, Homoptera Aphidoidea. 
Institut National de la Recerche Agronomique, 
Paris, 473 pp. 

Smith, C. E 1944. The Aphidiinae of North America 
(Braconidae: Hymenoptera). Ohio State Univer- 
sity Contributions in Zoology and Entomology 
No. 6, 154 pp. 

Stary, P. 1963. A study on the relationship of the My- 
zinae and their aphidiid parasites in (Central) Eu- 
rope (Homoptera: Aphidoidea, Myzinae; Hyme- 
noptera: Aphidiidae). Bolletino del Laboratorio di 
Entomologia Agraria “‘Filippo Silvestri” di Portici 
21:199-216. 

. 1966. Aphid Parasites of Czechoslovakia. W. 
Junk, The Hague, 242 pp. 

Stary, P. and C. FE Smith. 1976. The Nearctic distri- 
bution, host plants, and nomenclature of Monoc- 
tonus (Hymenoptera, Aphidiidae). Proceedings of 
the Entomological Society of Washington 78: 
171-175. 

van Achterberg, C. 1989. Revision of the subtribe 
Monoctonina Mackauer sensu stricto (Hymenop- 
tera: Braconidae: Aphidiinae). Zoologische Me- 
dedelingen 63: 1—22. 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 741-746 


ARCHIPS GOYERANA, N. SP. (LEPIDOPTERA: TORTRICIDAE) AN 
IMPORTANT PEST OF BALDCYPRESS (TAXODIACEAE) IN LOUISIANA 
AND MISSISSIPPI 


JAMES J. KRUSE 


Division of Insect Biology, Department of Environmental Science, Policy and Man- 
agement, University of California, Berkeley, CA 94720, U.S.A. (e-mail: kruse @nature. 
berkeley.edu) 


Abstract.—Archips goyerana, new species, is described and illustrated from southern 
Louisiana and southwestern Mississippi. It appears to be host-specific on baldcypress 
(Taxodium distichum; Taxodiaceae) and has become a serious pest of that tree species 
since its discovery in 1983. Notes on its biology are given, and differences between it 
and its sister species, A. argyrospila, are outlined. Although morphological differences 
between A. goyerana and A. argyrospila are subtle, they are corroborated by differences 


in forewing pattern, larval food plant preferences, and molecular data. 


Key words: 


Archips (Hiibner) is a large genus of at 
least 80 species widely distributed in the 
Nearctic, Palaearctic and Oriental regions 
(Razowski 1977, 1997). The fruittree leaf- 
roller, Archips argyrospila (Walker), is 
probably the most widespread of the ap- 
proximately 25 species of Archips in North 
America. It is a notorious, polyphagous pest 
of orchard trees throughout the northern 
United States and southern Canada (Chap- 
man and Lienk 1971, Smirle 1993, Meeker 
and Goyer 1994, Goyer and Chambers 
1997). Owing to the presence of numerous 
geographic forms and its considerable phe- 
notypic variation, MacKay (1962) and Goy- 
er et al. (1995) suggested that A. argyros- 
pila may consist of a number of different 
sibling species that are differentiated by 
food plant preference and/or geographical 
distribution. In this paper I describe A. goy- 
erana, a sibling species that feeds on bald- 
cypress (Taxodium distichum L. Rich., Tax- 
odiaceae) in Louisiana and Mississippi. 
This species has inflicted serious and wide- 


Tortricinae, Archipini, Taxodium 


spread defoliation in southern Louisiana 
since its discovery in 1983 (Braun et al. 
1990, Meeker and Goyer 1993, Goyer et al. 
1995). Aerial surveys conducted annually 
have found as many as 60,000 ha of bald- 
cypress forest affected by this insect in 
southern Louisiana (Goyer and Chambers 
1999): 


MATERIALS AND METHODS 


Institutions are abbreviated throughout 
the text as follows: Essig Museum of En- 
tomology (EMEC), University of Califor- 
nia, Berkeley, Ca, USA; Louisiana State 
Arthropod Museum (LSAM), Baton Rouge, 
LA, USA; Mississippi Entomological Mu- 
seum (MEM), Mississippi State, MS, USA; 
National Museum of Natural History 
(NMNH), Smithsonian Institution, Wash- 
ington. DC, USA; University of Minnesota, 
Saint Paul (UMSP), MN, USA. Specimens 
were obtained from the EMEC, MEM, and 
Louisiana State University Agricultural 
Center (LSUAC), Baton Rouge, LA, USA. 


742 


Dissection methodology follows that sum- 
marized in Brown and Powell (1991) ex- 
cept that preparations were transferred to 
95% isopropyl alcohol (instead of xylene) 
after the 95% EtOH wash, and all parts 
were slide-mounted with Euparol mounting 
medium (Bioquip, Gardena, CA) rather 
than Canada balsam. Forewing measure- 
ments were made with an ocular micro- 
meter mounted in a Zeiss Stemi SV6 dis- 
secting microscope. Terminology for geni- 
talic structures follows Horak (1984). Col- 
ors were described with the standards of 
Kornerup and Wanscher (1983). 


SYSTEMATICS 


Archips goyerana Kruse, new species 
(Figs. 1-2) 


Male (Fig. 1a).—Head: Vertex and up- 
per frons golden yellow to brownish or- 
ange, brown or brownish orange between 
antennae in most specimens. Labial palpus 
with golden yellow and brownish orange 
scaling. Mesonotum: Golden yellow and 
brownish orange scaling, dark reddish 
brown scaling also present in many speci- 
mens; tegulae usually concolorous with me- 
sonotum, but often with more brownish or- 
ange or reddish brown scaling. Forewing: 
Length 6.8-8.5 mm (mean 7.4 mm, n = 
37). Costal fold extending from base to near 
proximal margin of median fascia, i.e., ca. 
30-35% length of FW costa. Upper side 
with underscaling gray to grayish red, gray- 
ish red concentrated near center, extending 
basally and dorsally; overscaling pattern el- 
ements include distinct but broken reddish 
brown fasciae and black, broken strigulae 
and irregular white, yellowish white, gray, 
and golden yellow scaling throughout; bas- 
al fasciae indistinct; subbasal and median 
fasciae distinct and strongly contrasting 
with yellowish white interfascial regions 
near costa; postmedian fasciae distinct at 
costa, breaking dorsally; subterminal and 
terminal fasciae indistinct, represented by 
reddish brown strigulae or reddish brown 
patches, broken by yellowish white and 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


golden yellow scaling. Fringe gray with 
reddish brown and/or brownish orange 
scales replaced by gray scales at tornus. 
Under side with upper side pattern near cos- 
ta, becoming gray dorsally in area of 
hindwing overlap. Hindwing: Upper side 
uniformly gray, except some dark gray and/ 
or orange scaling near apex in some indi- 
viduals. Under side gray, yellowish white 
near costa, orange strigulae or dark gray or 
orange scaling near apex. Genitalia: As in 
Fig. 1b (slide JJK 220; n = 6). Uncus large, 
prominent, slightly narrowed from base, 
slightly broadened apically, width of apex 
generally less than or equal to base (Fig. 
lc). Gnathos arising from triangular projec- 
tions of the tegumen, gently concavely 
curved and fused together apically, extend- 
ing slightly beyond coastal margin of valva; 
socius vestigal; valva ovate; sacculus grad- 
ually broadening terminad with produced 
tip arising subapically of valva. Aedeagus 
straight or gently curved, terminating in 
slender tip strongly curved to left. Phallo- 
base gently curved (Fig. 1d). 

Female (Fig. 2a).—FW length 8.0—10.0 
mm (mean 8.9 mm, n = 50). Superficially 
as in male except forewing ground color 
gray, with very little grayish red near center 
of wing, and generally with less yellowish 
white and golden yellow overscaling, but 
more black strigulae throughout. Genitalia: 
As in Fig. 2b (slide JJK 221; n = 7). Pa- 
pillae anales setose, without projections; 
sterigma short, produced as a shallow bowl; 
antrum ovate; ductus bursae long; signa 
long, bladelike. Genitalia are virtually in- 
distinguishable from A. argyrospila except 
for a more weakly produced sterigma (Fig. 
2G). 

Types.—Holotype ¢: LOUISIANA: As- 
sumption Parish, Pierre Part, 27 April 1999 
as pupa (R. A. Goyer), in LSAM. Paraty- 
pes: (n = 101). LOUISIANA: Assumption 
Parish, Pierre Part, 27 April 1999 as pupae, 
14 3, 23 2 (R. A. Goyer); St. Charles Par- 
ish, Norco, 27 April 1999 as pupae, 19 6, 
42 2 (R. A. Goyer); MISSISSIPPI: Hinds 
County, Clinton, 2 2, 18-19 May 1996 (M. 


VOLUME 102, NUMBER 3 743 


( L a : 


Figs. 1-2. Adults and genitalia of Archips goyerana. la, Male of A. goyerana, holotype, Assumption Parish, 
Louisiana; scale in mm. 1b, Male of A. goyerana; valvae spread, aedeagus removed, holotype, Assumption 
Parish, Louisiana, slide number JJK 220.; scale bar = 1 mm. Ic, Detail of uncus of male A. goyerana (left) and 
detail of uncus of male A. argyrospila (right), Harrison County, Long Beach, Mississippi, drawn from slide 
number JJK 117. 1d, Detail of aedeagus of male A. goyerana (left) and detail of aedeagus of male A. argyrospila 
(right). 2a, Female of A. goyerana, paratype, St. Charles Parish, Louisiana. scale in mm. 2b, Female of A. 
goyerana, paratype, St. Charles Parish, Louisiana, slide number JJK 221; scale bar = 1 mm. 2c, Detail of 
sterigma of female A. goyerana (left) and detail of sterigma of female A. argyrospila (right), Bossier Parish, 
Barksdale Air Force Base, Louisiana, drawn from slide number JJK 214. 


and E. Roshore); Harrison County, Long Diagnosis.—Archips goyerana was dis- 
Beach, | 2, 13 May 1997 (R. Kergosien). covered in 1983 in Iberville Parish, Loui- 
Paratypes are deposited in LSAM (36), | siana (Goyer and Lenhard 1988). It is very 
MEM (10), UMSP (5), NMNH (20), and — similar in appearance to A. argyrospila but 
EMEC (30). it is far less variable. In A. goyerana the 


744 


fasciae are usually dark reddish brown, in- 
terfascial spots are strongly contrasting with 
fasciae, and black striae are usually present. 
Interfascial spots are generally less well de- 
fined in A. argyrospila, and fasciae typical- 
ly consist of various shades of brown or 
brownish orange. A pale form of A. goy- 
erana is brownish orange on the forewing 
and pale brownish orange or light gray on 
the hindwing. This form appears to domi- 
nate in Mississippi and is difficult to distin- 
guish from co-occurring A. argyrospila. In 
male genitalia of A. argyrospila the uncus 
is distinctly widened terminally, weakly 
club shaped (Fig. Ic). In addition, the phal- 
lobase in A. argyrospila is typically curved 
at an angle greater than 45° from the plane 
of the aedeagus as opposed to less than 35° 
in A. goyerana (Fig. 1d). Female genitalia 
of A. goyerana are nearly indistinguishable 
from those of A. argyrospila, although the 
sterigma tends to be slightly more robust 
and squarish in A. argyrospila, in compar- 
ison to the shallow mildly angled bowl of 
A. goyerana (Fig. 2c). Molecular data in- 
dicates a divergence ranging from 1.47 to 
2.53% between A. argyrospila and A. goy- 
erana in 475 base pairs of mitochondrial 
DNA in the Cytochrome Oxydase I gene, 
while widely separated populations of A. 
argyrospila differ by 1.26% or less (Kruse 
and Sperling, unpublished data). 
Distribution and biology.—Archips goy- 
erana occurs in southern Louisiana and 
southwestern Mississippi (Fig. 3). It origi- 
nally may have been endemic to forested 
wetlands, but has undergone a population 
explosion and expansion of its range since 
1983 (Goyer et al. 1990, Zhou et al. 1993, 
Goyer et al. 1995). In 1988, A. goyerana 
was found for the first time on baldcypress 
east of the Mississippi River (in Baton 
Rouge) and by 1993 had spread eastward 
to the suburbs of New Orleans (Goyer et al. 
1995). Noticeable defoliation was evident 
Over portions of at least eleven Louisiana 
parishes (Meeker and Goyer 1993, 1994). 
It was collected in southwestern Mississippi 
in 1996 (Clinton) and in 1997 (Long 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Beach). Archips argyrospila and A. goyer- 
ana are known to co-occur near Baton 
Rouge, Louisiana and Long Beach, Missis- 
sippi. Individuals captured away from for- 
ested wetlands are speculated to feed on or- 
namental baldcypress. 

Like A. argyrospila, A. goyerana 1s uni- 
voltine throughout its range, with overwin- 
tering egg masses attached to the bark of 
thin twigs of the host plant in obligate dia- 
pause (Braun et al. 1990, Goyer et al. 1990, 
Brown 1991, Meeker and Goyer 1993, 
Goyer and Chambers 1997). Dormant eggs 
hatch by bud break of baldcypress trees 
during late February and early March (Goy- 
er et al. 1990, Meeker and Goyer 1993, 
Goyer and Chambers 1997). First instar lar- 
vae disperse and seek out the terminal por- 
tions of expanding baldcypress foliage, bur- 
row within the cluster of young needles, 
and begin feeding inconspicuously (Goyer 
and Chambers 1997). 

Larvae of A. goyerana feed on the open- 
ing leaf buds and elongating leaves (Braun 
et al. 1990). As the foliage expands, devel- 
oping A. goyerana larvae produce silk to 
roll adjacent needles and branchlets into a 
tight mass, surrounding themselves individ- 
ually and then feeding on the foliage inside 
(Braun et al. 1990, Goyer and Chambers 
1997). Larvae remain inconspicuous on 
baldcypress unless the tree is severely de- 
foliated, at which time they become very 
active, crawling on branches and spinning 
down on silk (Braun et al. 1990). Massive 
mortality to mature larvae by drowning in 
standing water was observed on many oc- 
casions where baldcypress were completely 
defoliated (Braun et al. 1990). 

Larvae undergo five larval instars before 
pupation (Braun et al. 1990). Adults emerge 
between late-April and mid-May (Goyer 
and Chambers 1997). The development 
time for A. argyrospila from egg hatch to 
adult emergence takes about 6 weeks 
(Braun et al. 1990). For A. goyerana, de- 
velopment time is considerably longer, 
about 8 to 10 weeks on baldcypress in the 
field (Braun et al. 1990, Goyer and Cham- 


VOLUME 102, NUMBER 3 


745 


Fig. 3. 
occur are shaded. 


bers 1997). Previous studies have shown 
that A. goyerana is virtually unable to sur- 
vive on oaks, a major food source for near- 
by populations of A. argyrospila (Goyer et 
al. 1995). Differences in pheromonal re- 
sponses between citrus-feeding A. argyros- 
pila and baldcypress-feeding A. goyerana 
populations suggest evolutionary diver- 
gence in mating behavior (Goyer et al. 
1995). The forested wetland habitat of A. 
goyerana and the distant relationship be- 
tween its food plant and the food plants of 
A. argyrospila further confirm that the two 
are distinct. 

Archips goyerana is a wetlands pest in 
Louisiana. Repeated defoliation of baldcy- 


Range of Archips goyerana in Louisiana and Mississippi. Counties where A. goyerana is known to 


press has been observed in several areas of 
southern Louisiana, with dramatic reduc- 
tions in radial growth and death of portions 
of the tree canopy in all age groups, and 
mortality primarily in the small, understory 
trees that do not recover fully from com- 
plete and repeated defoliation (Goyer and 
Lenhard 1988, Goyer et al. 1990, Goyer 
and Chambers 1997). In addition, baldcy- 
press defoliation is closely linked with ex- 
cessive flooding, affecting tree successional 
patterns in forest wetlands (Goyer and 
Chambers 1997). With 70% of Louisiana’s 
baldcypress growing stock located within or 
near areas now infested with A. goyerana, 
the persistence and continued spread of de- 


746 


foliation poses an additional threat to this 
dwindling and unique forest resource 
(Meeker and Goyer 1993). 

Etymology.—The species is named for 
Richard A. Goyer of the Department of En- 
tomology, Louisiana Agricultural Experi- 
ment Station, Louisiana State University 
Agricultural Center, Baton Rouge, Louisi- 
ana. He collected the holotype as well as 
most of the paratypes, and is responsible for 
the majority of the research on the biology 
and ecology of the species. 


ACKNOWLEDGMENTS 


I thank Richard A. Goyer (LSUAC) and 
Richard L. Brown (MEM) for the contri- 
bution of specimens, and the collectors cit- 
ed in the text, who made this study possi- 
ble. I thank the following for reviewing and 
providing helpful comments on various 
drafts of the manuscript: John W. Brown, 
Systematic Entomology Laboratory, 
USDA, Washington, DC; Jerry A. Powell, 
University of California, Berkeley, CA; 
Brent D. Mishler, University of California, 
Berkeley, CA; and Felix A. H. Sperling, 
University of Alberta, Edmonton, Alberta. 
This work was supported by a National Sci- 
ence Foundation Partnerships in Enhancing 
Expertise in Taxonomy (PEET) grant to Fe- 
lix A. H. Sperling and Jerry A. Powell. 


LITERATURE CITED 


Braun, D. M., R. A. Goyer, and G. J. Lenhard. 1990. 
Biology and mortality agents of the fruit tree leaf- 
roller (Lepidoptera: Tortricidae), on baldcypress in 
Louisiana (USA). Journal of Entomological Sci- 
ence 25: 176-184. 

Brown, J. J. 1991. Diapause, pp. 175—185. Jn vari der 
Geest, L. P. S. and H. H. Evanhuis ,eds., Tortricid 
Pests: Their Biology, Natural Enemies and Con- 
trol. World Crop Pests,. Vol. 5. Elsevier, Holland. 
808 pp. 

Brown, J. W. and J. A. Powell. 1991. Systematics of 
the Chrysoxena group of genera Lepidoptera: Tor- 
tricidae: Euliini). University of California Publi- 
cations in Entomology 111, 87 pp. 

Chapman, P. J. and S. E. Lienk. 1971. Tortricid fauna 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


of apples in New York (Lepidoptera: Tortricidae). 
New York State Agricultural Experiment Station 
Special Publication, 122 pp. 

Goyer, R. A. and J. L. Chambers. 1997. Evaluation of 
insect defoliation in baldcypress and its relation- 
ship to flooding. National Biological Service Bi- 
ological Science Report 8, 36 pp. 

Goyer, R. A. and G. J. Lenhard. 1988. A new insect 
pest threatens baldcypress. Louisiana Agricultural 
Experiment Station Bulletin 31: 16—21. 

Goyer, R. A., G. J. Lenhard, and J. D. Smith. 1990. 
Insect herbivores of a baldcypress-tupelo ecosys- 
tem. Forest Ecology and Management 33-34: 
517-522. 

Goyer, R. A., T. D. Paine, D. P. Pashley, G. J. Lenhard, 
J. R. Meeker, and C. C. Hanlon. 1995. Geographic 
and host-associated differentiation in the fruittree 
leafroller (Lepidoptera: Tortricidae). Annals of the 
Entomological Society of America 88: 391-396. 

Horak, M. 1984. Assessment of taxonomically signif- 
icant structures in Tortricinae (Lep., Tortricidae). 
Bulletin de la Société Entomologique Suisse 57: 
3-64. 

Kornerup, A. and J. H. Wanscher. 1983. Methuen 
handbook of colour (3rd ed.). Methuen and Com- 
pany, London. 

MacKay, M. R. 1962. Larvae of the North American 
Tortricinae (Lepidoptera: Tortricidae). Canadian 
Entomologist Supplement 28: 1—182. 

Meeker, J. R. and R. A. Goyer. 1993. Relationships 
between patterns of defoliation by the fruittree 
leafroller (Lepidoptera: Tortricidae) and foliage 
morphology of baldcypress in forested wetlands 
of Louisiana. Journal of Entomological Science 
28: 317-326. 

. 1994. Fruittree leafroller, Archips argyrospila 
(Walker) (Lepidoptera: Tortricidae), performance 
as influenced by host foliage type. Journal of En- 
tomological Science 29: 1-9. 

Razowski, J. 1977. Monograph of the genus Archips 
Hiibner (Lepidoptera, Tortricidae). Actazoologica 
Cracoviensia 22: 55—206, 223. 

. 1997. Generic composition of the New World 
Archipini (Lepidoptera, Tortricidae) with descrip- 
tion of two new genera and two new species. Mis- 
cel-lania Zoologica 20.1: 125-130. 

Smirle, M. J. 1993. Larval performance of two leaf- 
roller species on known and potential hosts. En- 
tomologia Experimentalis et Applicata 67: 223-— 
Bil. 

Zhou, J., R. A. Goyer, and G. J. Lenhard. 1993. Sea- 
sonal occurrence of Calleida viridipennis Say and 
Plonchionus timidus Haldeman (Coleoptera: Car- 
abidae) predators of the fruit tree leafroller Ar- 
chips argyrospila Walker in Louisiana. Journal of 
Entomological Science 28: 254—262. 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 747-754 


DISCOVERY OF THE SMALL JAPANESE CEDAR LONGHORNED BEETLE, 
CALLIDIELLUM RUFIPENNE (MOTSCHULSKY) 
(COLEOPTERA: CERAMBYCIDAE), IN LIVE ARBORVITAE 
IN CONNECTICUT 


CHRIS T. MAIER AND CAROL R. LEMMON 


Department of Entomology, Connecticut Agricultural Experiment Station, PO. Box 
1106, New Haven, CT 06504, U.S.A. (e-mail: chris.maier@po.state.ct.us) 


Abstract.—The small Japanese cedar longhorned beetle, Callidiellum rufipenne (Mot- 
schulsky) (Coleoptera: Cerambycidae), was discovered principally in three cultivars of 
American arborvitae, Thuja occidentalis L., in four towns in Fairfield and New Haven 
Co. in southwestern Connecticut. Between September 1998 and March 1999, infestations 
of C. rufipenne were found in 102 (0.5%) of 20,000 cupressaceous plants that were 
inspected in garden centers, nurseries, landscaped areas, and the wild in Connecticut. 
Within 2 years of planting, 94 (92.2%) of infested plants had been balled and burlaped, 
which may have stressed them. Of the 102 plants, 101 (99%) were 0.9-2.1 m high ar- 
borvitae that averaged 6.3 beetles/plant, and the other one was yellow cedar, Chamae- 
cyparis nootkatensis (D. Don) Sprach. Arrivals of beetles in wood material at ports of 
entry, increased international trade and travel, and the high abundance of potential hosts 
in coastal North America probably have facilitated introduction and establishment. Adults, 


life cycle, and damage of C. rufipenne are described briefly. 


Key Words: 


Arborvitae, Callidiellum, Cerambycidae, Chamaecyparis, Coleoptera, Con- 


necticut, Cupressaceae, Juniperus, larval damage, plant pest, state record, 


Thuja occidentalis 


The small Japanese cedar longhorned 
beetle, Callidiellum rufipenne (Motschul- 
sky) (Coleoptera: Cerambycidae), is best 
known as a borer of Japanese cedar, Cryp- 
tomeria japonica D. Don (Taxodiaceae), 
and Japanese cypress or Hinoki false cy- 
press, Chamaecyparis obtusa (Siebold & 
Zuccarini) Endlicher (Cupressaceae) (Ma- 
kihara 1984, Shibata 1994), which are im- 
portant trees in Japanese plantations (Ko- 
bayashi 1985). Larvae of C. rufipenne, 
however, also tunnel into the wood of other 
species of Cupressaceae and rarely Pina- 
ceae (Bates 1873, Gressitt 1951, Shiraki 
1952, Campadelli and Sama 1988, Bahillo 
and Iturrondobeitia 1995). Shibata (1994) 


and others have suggested that C. rufipenne 
attacks only dead or dying wood of its co- 
niferous hosts. Other species of Callidiel- 
lum, C. cupressi (Van Dyke) and C. vires- 
cens Chemsak and Linsley in the western 
United States (Chemsak and Powell 1964, 
Chemsak and Linsley 1966) and C. flavo- 
signatus Pu and C. villosulum (Fairmaire) 
in China (Gressitt 1951, Pu 1991), also are 
restricted to conifers in the Cupressaceae, 
Pinaceae, and Taxodiaceae. 

In May 1997, a single adult of C. rufi- 
penne was collected on wild eastern red ce- 
dar, Juniperus virginiana L., at Manteo, 
Dare Co., North Carolina (E.R. Hoebeke, in 
litt.). This represented the first record of this 


748 


eastern Asian beetle in the wild in the Unit- 
ed States. After the initial discovery of the 
lone adult, K. Ahlstrom (in litt.) reared ad- 
ditional adults from larvae collected in the 
wood of dead eastern red cedar at Manteo. 
Before its recent introduction into Italy 
(Campadelli and Sama 1988), Spain (Bah- 
illo and Iturrondobeitia 1995), and this 
country, C. rufipenne had a distributional 
range of Russia (Sakhalin), Japan (includ- 
ing the Ryukyu Islands), Korea, Taiwan, 
and northeastern China (e.g., Gressitt 1951, 
Duffy 1968, Makihara et al. 1989). Gressitt 
(1951) suggested that C. rufipenne probably 
was introduced into Taiwan. Port inspectors 
have repeatedly intercepted this beetle in 
dunnage and other wood products shipped 
to New Zealand (Bain 1974, 1977), the 
United States (Mumford 1965, 1966, 1967; 
Girard) 1968; 1969; 119715,.1972a;-01972b, 
1973, 1974; USDA 1979, 1980, 1981, 
1982, 1984), and other countries. 

Here we report the discovery of C. rufi- 
penne in Connecticut, document its infes- 
tation of live plants, mainly American ar- 
borvitae or northern white cedar, Thuja oc- 
cidentalis L., and briefly describe its life 
history and the appearance of damage. We 
initiated this and additional studies because 
C. rufipenne may pose a threat to the nurs- 
ery industry and to northern forests. Arbor- 
vitae are important to the nursery trade be- 
cause they are popular ornamental shrubs 
grown widely and planted especially near 
homes and commercial buildings. In addi- 
tion, northern white cedars are an integral 
part of some forests in northeastern and 
north-central North America. 


MATERIALS AND METHODS 


Adult records from Connecticut are 
based on inspecting plants for larval injury 
and then removing or rearing beetles from 
plants with damage. We extracted adults 
from wood mainly between late September 
1998 and February 1999 by finding holes 
(the eventual adult exit routes that had been 
plugged with chewed wood by larvae) in 
branches and then by splitting the wood to 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


reveal the individual adults in their pupal 
chambers. Between November 1998 and 
March 1999, additional infested arborvitae 
(about 80% of ‘all infested plants) were 
transported from garden centers or land- 
scaped areas, mostly residential yards in 
Fairfield and New Haven Co., to two out- 
door enclosures with double screening at 
Lockwood Farm, Hamden, New Haven 
Co., Connecticut. In late March, branches 
that appeared to be uninfested were pruned 
from these shrubs and burned in accordance 
with state regulations. The remaining wood 
that had evidence of larval boring was di- 
vided into two equal groups. Wood of each 
of the shrubs in one lot was placed in a 
sealed white plastic drum (18.9 liters) in an 
environmental chamber adjusted to 21 + 
1°C and to a 15L:9D photoperiod. The re- 
mainder of the wood was put into screened 
cages of various sizes in one of the outdoor 
screened houses. Adults were collected 
from drums or cages every 3—7 days until 
emergence ended. 

Data for infestations of C. rufipenne dis- 
covered by federal inspectors at ports of en- 
try between 1964 and 1982 (Mumford 
1965, 1966, 1967; Girard 1968, 1969, 1971, 
1972a, 19726, 1973, 1974; USDASIC7= 
1980, 1981, 1982, 1984) were used in Fig. 
1. Yearly records were based on the fiscal 
year, which changed during the mid-1970’s. 
Between 1964 and 1982, C. rufipenne had 
a quarantine status of an actionable pest. In 
1982, the Animal and Plant Health Inspec- 
tion Service (APHIS) changed its quaran- 
tine status from an actionable to a non-re- 
portable pest (J. Cavey, in litt.), which re- 
sulted in a rapid decrease in records of its 
interception. In 1998, C. rufipenne again 
became an actionable pest. 

Voucher specimens are deposited at the 
Connecticut Agricultural Experiment Sta- 
tion, New Haven; Cornell University, Itha- 
ca, New York; National Museum of Natural 
History, Smithsonian Institution, Washing- 
ton, D.C.; and Essig Museum of Entomol- 
ogy, University of California, Berkeley, 
California. 


VOLUME 102, NUMBER 3 


Table 1. 


749 


Location and characteristics of live arborvitae, Thuja occidentalis, infested by the small Japanese 


cedar longhorned beetle, Callidiellum rufipenne, in Connecticut during 1998. 


Location of Infested Plants No. Beetles 

County Town Variety of Arborvitae Height (m) No. Infested Mean Range 
Fairfield Greenwich Smaragd 1.8—2.1 7 4.4 2-9 
Fairfield Greenwich/Stamford! Smaragd 1.5-1.8 4 10.3 2-27 
Fairfield Stamford Brandon 1.8—2.1 2 4.0 3-5 
Fairfield Stamford Smaragd 1.5-1.8 26 Hitt 2-37 
Fairfield Stamford/Greenwich! Smaragd 1.5-1.8 16 S55) 2-11 
New Haven Milford Nigra 1.5-1.8 2 2.0 
New Haven Milford Smaragd 0.9-1.5 8 IBS 1-3 
New Haven Milford Smaragd 1.2-1.8 Si 4.5 1-16 
New Haven North Haven Smaragd 1.5-1.8 5 3.4 1-9 


'in 1998, these arborvitae initially were at the garden center in the town listed first, and then, usually in 
spring of the same year, they were planted in yards in the town listed second. 


RESULTS AND DISCUSSION 


Infestations in Connecticut.—On 22 Sep- 
tember 1998, one adult of C. rufipenne was 
found in a branch of a emerald green ar- 
borvitae, T. occidentalis ‘Smaragd’, planted 
in spring 1998 near a small factory in Mil- 
ford, New Haven Co. Subsequent inspec- 
tion of the planting of 42 arborvitae at this 
site revealed 7 live plants (16.7% of the to- 
tal) with injury from larval boring. The 42 
arborvitae planted by the factory were pur- 
chased at a nearby garden center that also 
had infested arborvitae that had not been 
sold. This find represents the second North 
American record of this beetle outside a 
port of entry, and the first from a live ar- 
borvitae that visually appeared to be 
healthy. In 1967, an inspector at a port in 
the United States did intercept C. rufipenne 
in dunnage of a Thuja species on a cargo 
ship (Girard 1968). 

Between autumn 1998 and spring 1999, 
state inspectors found 102 (0.5% of total) 
infested plants among 20,000 cupressa- 
ceous shrubs or trees examined in garden 
centers, nursery fields, landscaped areas, 
and the wild throughout Connecticut. Plants 
infested with C. rufipenne were located 
mainly in four garden centers and in nearby 
residential areas with arborvitae that re- 
cently had been purchased from the garden 
centers (see Table 1 for the locations). Of 


102 infested plants, 100 (98.0%) had been 
imported from western North America, and 
94 (92.2%) had been balled and burlaped 
within 2 years of inspection for C. rufipen- 
ne. Thorough examination of nursery fields 
at the points of origin in British Columbia 
(J. Bell and B. Gill, in litt.) and Oregon (R. 
Westcott, in litt.) and inspection of ship- 
ments from the western North America in 
1999 did not reveal any damage or speci- 
mens of C. rufipenne. The plants with C. 
rufipenne apparently became infested in 
Connecticut; however, the origin and time 
of introduction are unknown. 

Of the 102 infested plants, 101 (99.0%) 
were arborvitae (Table 1), and one was a 
yellow cedar, Chamaecyparis nootkatensis 
(D. Don) Spach. Among the 101 infested 
arborvitae of three cultivars, 97 (96.0%) 
were the variety ‘Smaragd’, which current- 
ly is the most widely grown and planted 
arborvitae in Connecticut. Infested arbor- 
vitae had between 1 and 37 beetles, and 
ranged in height from 0.9 to 2.1 m. In all, 
632 beetles developed in the wood of the 
101 arborvitae (mean = 6.3 live beetles/ 
plant), and 6 in that of yellow cedar. An 
unknown number of arborvitae that may 
have been infested by C. rufipenne at the 
four garden centers were sold and probably 
planted in southwestern Connecticut. 

Even though most infested plants that 


750 


100 


No. Interceptions 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


64 66 68 70 72 74 76 78 80 82 


Fiscal Year 


Figaale 


Frequency of interceptions of the small Japanese cedar longhorned beetle, Callidiellum rufipenne, at 


ports of entry in the United States between 1964 and 1982. The change in the fiscal year in the early 1970's 
apparently caused some inconsistencies in compilation of data between 1973 and 1975. 


were balled and burlaped appeared healthy, 
they may have been susceptible to attack by 
C. rufipenne because they were stressed by 
root and branch breakage and possibly in- 
adequate watering before, during, and after 
shipment to Connecticut. Furthermore, 
these arborvitae occasionally were infested 
by bark beetles (Scolytidae), such as 
Phloeosinus canadensis Swaine and Poly- 
graphus rufipennis (Kirby). 

Factors contributing to accidental intro- 
duction and establishment.—The small Jap- 
anese cedar longhorned beetle has repeat- 
edly been intercepted by federal and state 
inspectors at ports of entry in the United 
States (Mumford 1965, 1966, 1967; Girard 
19685 1969, 197 13 197 2a 19726; 6 1973; 
1974; “OSDA 11979, 1980) 19811982, 
1984; R. Penrose, in litt.). In most cases, 
the intercepted beetles were associated with 


dunnage or other wooden material from 
eastern Asia. In Vancouver, British Colum- 
bia, in 1927, C. rufipenne was reared from 
imported wood of Japanese origin (Leech 
1949). Hatch (1971) recorded another spec- 
imen from Seattle, Washington, but this 
beetle, too, was likely associated with an 
imported wood product. 

Between 1964 and 1982, when federal 
inspectors actively reported infestations of 
C. rufipenne, they repeatedly discovered 
them in imported wood material (Fig. 1). 
During these years, infestations were found 
in dead wood, especially dunnage, in 21 
states in the United States and in Puerto 
Rico (Mumford 1965, 1966, 1967; Girard 
1968, 1969, 1971; 1972a,. 1972b; 799723 
1974; USDA 1979, 1980; 198171982; 
1984). Although C. rufipenne was not dis- 
covered at ports of entry in Connecticut, it 


VOLUME 102, NUMBER 3 


was found in ports of the nearby states of 
New Jersey, New York, and Pennsylvania. 
In coastal counties in Connecticut and near- 
by states, potential hosts abound in residen- 
tial areas and forests. In particular, over- 
mature, often dead and dying trees of J. vir- 
giniana are numerous in successional areas 
along major highways and in regrowth for- 
ests. We suggest that the frequency of ar- 
rivals of C. rufipenne in imported wood, in- 
creased international trade and travel, and 
the high abundance of potential hosts near 
coastal areas probably have facilitated the 
accidental introduction and establishment 
of C. rufipenne in the United States. 

Brief description of adult appearance, life 
history, and damage.—Males are iridescent 
deep blue with a single brownish red to red 
patch at the anterolateral corner of the up- 
per surface of each elytron. The females 
(Fig. 2) have entirely reddish brown elytra 
and a reddish abdomen. The antennae of 
males are longer than the body, whereas 
those of females are shorter, about % to % 
the length of the body. Thus, the sexes can 
be separated by both color and antennal 
length. When they are reared from arbor- 
vitae, adults vary from 5—13 mm in length. 

The life history of C. rufipenne (Shibata 
1994, in litt.; Maier 1999; Y. Soma, in litt.) 
is similar to that of other species of Calli- 
diellum (Chemsak and Powell 1964, Nak- 
amura and Kojima 1981). Adults of C. ru- 
fipenne emerge from wood in spring and 
soon mate on the host. Females of C. rufi- 
penne then lay eggs singly or in small 
groups in cracks or crevices in stems or 
bark. After hatching, larvae bore into the 
cambium and phloem. They expand their 
feeding tunnel until they become full-grown 
(Fig. 3) in late summer. After feeding is 
completed, larvae bore into the sapwood 
and carve an ellipsoidal pupal cell. The pu- 
pal chamber is connected to the surface of 
a branch by an exit tunnel (Fig. 4) that is 
oval in cross-section. Larvae plug the even- 
tual exit route with fragments of chewed 
wood. They pupate in late summer or early 
fall, and adults eclose within 2—3 weeks, 


ial 


usually in the autumn. Adults remain in the 
pupal cell until the following spring when 
they remove the plug to the pupal cell and 
emerge. In Connecticut, C. rufipenne has 
one generation per year. In Japan, the life 
cycle also usually requires 1 year. In some 
northern regions of Japan, however, it may 
require 2 years to complete the life cycle 
(Y. Soma, in litt.). 

In live arborvitae, injury from larval bor- 
ing becomes most obvious between late 
summer and the following spring when the 
bark of small arborvitae often splits to re- 
veal the sinuous larval tunnels (Fig. 5). The 
larval gallery is packed with frass, which 
apparently is also mixed with chewed wood 
fragments. In some cases, these tunnels may 
encircle branches and, thereby, disrupt the 
flow of water and nutrients within the plant. 
The material packed in the larval gallery 
has both light and dark particles, giving it 
a “‘salt-and-pepper”’ look. 


ACKNOWLEDGMENTS 


We thank Xavier Asbridge, Joan Bravo, 
Dora Brown, Jeffrey Fengler, Verryn Jen- 
nings, Tracy LaProvidenza, Roland Leval- 
lee, Gale Ridge-O’Connor, Stephen San- 
drey, Michael Thomas, and Peter Trenchard 
for assistance in inspecting plants and rear- 
ing beetles. We especially want to acknowl- 
edge Gale Ridge-O’Connor, who recog- 
nized that the beetle was not native to Con- 
necticut. The screened buildings for storing 
infested plants were constructed by Richard 
Cecarelli, Jeffrey Fengler, Rollin Hannan, 
and Edward Naughton. Adults of C. rufi- 
penne were identified by John A. Chemsak, 
University of California, Berkeley, E. Ri- 
chard Hoebeke, Cornell University, Ithaca, 
and Steven A. Lingafelter, USDA System- 
atic Entomology Laboratory, Washington, 
D.C. Donald E. Bright, Jr., Research 
Branch, Agriculture and Agri-Food Canada, 
Ottawa, Ontario, identified the scolytids. 
Kenneth Ahlstrom, North Carolina Depart- 
ment of Agriculture, Raleigh, Jon Bell, Ca- 
nadian Food Inspection Agency, New West- 
minster, British Columbia, Joseph Cavey, 


752 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 2-5. Small Japanese cedar longhorned beetle, Callidiellum rufipenne. 2, Adult female, 12 mm in length. 
3, Full-grown larva exposed after removal of bark of arborvitae, 13 mm in length. 4, Larva in its ellipsoidal 
pupal cell; note the pupal plug of chewed wood fragments. 5, Branch injury caused by larval boring. Arrows 
show frass and chewed wood exposed after the bark splits in late summer. 


VOLUME 102, NUMBER 3 


USDA, APHIS, PPQ, Riverdale, Maryland, 
Bruce Gill, Canadian Food Inspection 
Agency, Ottawa, Ontario, E. Richard Hoe- 
beke, Richard Penrose, California Depart- 
ment of Food and Agriculture, Sacramento, 
Eiichi Shibata, Nagoya University, Japan, 
Yukihiro Soma, Yokohama Plant Protection 
Station, Yokohama, Japan, and Richard 
Westcott, Oregon Department of Agricul- 
ture, Salem, provided valuable information 
about C. rufipenne or surveys for it. Robert 
Tracy, USDA, APHIS, PPQ, Elizabeth, 
New Jersey, kindly provided access to pub- 
lications that could not easily be obtained 
elsewhere. Finally, we appreciate the co- 
operation of Patricia Douglass and Wayne 
Petrie, USDA, APHIS, PPQ, Wallingford, 
Connecticut. 


LITERATURE CITED 


Bahillo, P. and J. C. Iturrondobeitia. 1995. Primera cita 
de Callidiellum rufipenne (Motschulsky, 1860) 
para la Peninsula Ibérica (Coleoptera: Ceramby- 
cidae). Boletin de la Asociaci6n Espanola de En- 
tomologica 19(3—4): 204. 

Bain, J. 1974. Overseas wood- and bark-boring insects 
intercepted at New Zealand ports. New Zealand 
Forest Service (Wellington) Technical Paper No. 
61, 24 pp. 

. 1977. Overseas wood- and bark-boring insects 
intercepted at New Zealand ports. New Zealand 
Forest Service (Wellington) Technical Paper No. 
63, 28 pp. 

Bates, H. W. 1873. On the longicorn Coleoptera of 
Japan. Annals and Magazine of Natural History, 
including Zoology, Botany, and Geology (4™ se- 
ries) 12: 193—201. 

Campadelli, G. and'G. Sama. 1988. Prima segnalazio- 
ne per |’Italia di un cerambicide giapponese: Cal- 
lidiellum rufipenne Motschulsky. Bollettino 
dell’Istituto di Entomologia «Guido Grandi» 
dell’ Universita di Bologna 43: 69-73. 

Chemsak, J. A. and E. G. Linsley. 1966. The genus 
Callidiellum in North America (Coleoptera: Cer- 
ambycidae). Pan-Pacific Entomologist 42: 151— 
1S5i, 

Chemsak, J. A. and J. A. Powell. 1964. Observations 
on the larval habits of some Callidiini with special 
reference to Callidiellum cupressi (Van Dyke) 
(Coleoptera: Cerambycidae). Journal of the Kan- 
sas Entomological Society 37: 119-122. 

Duffy, E. A. J. 1968. A Monograph of the Immature 
Stages of Oriental Timber Beetles (Cerambyci- 


753 


dae). The British Museum (Natural History), Lon- 
don, England, 434 pp. 

Girard, D. H., compiler. 1968. List of intercepted plant 
pests, 1967 (pests recorded from July 1, 1966, 
through June 30, 1967). United States Department 
of Agriculture, Agricultural Research Service, 
Plant Quarantine Division, Hyattsville, Maryland. 
ARS 82-6-2, 76 pp. 

. 1969. List of intercepted plant pests, 1968 

(pests recorded from July 1, 1967, through June 

30. 1968). United States Department of Agricul- 

ture, Agricultural Research Service, Plant Quar- 

antine Division, Hyattsville, Maryland. ARS 82- 

6-3, 86 pp. 

. 1971. List of intercepted plant pests, 1969 

(pests recorded from July 1, 1968, through June 

30, 1969). United States Department of Agricul- 

ture, Agricultural Research Service, Plant Quar- 

antine Division, Hyattsville, Maryland. ARS 82- 

6-4, 77 pp. 

. 1972a. List of intercepted plant pests, 1970 

(pests recorded from July 1, 1969, through June 

30, 1970). United States Department of Agricul- 

ture, Animal and Plant Health Inspection Service, 

Hyattsville, Maryland. APHIS 82-1, 85 pp. 

. 1972b. Addendum to list of intercepted plant 

pests, 1970 (pests recorded from July 1, 1969, 

through June 30, 1970). United States Department 

of Agriculture, Animal and Plant Health Inspec- 

tion Service, Hyattsville, Maryland. APHIS 82-1 

(we suspect that this issue should have been la- 

beled 82—2), 13 pp. 

. 1973. List of intercepted plant pests, 1971 

(pests recorded from July 1, 1970, through June 

30, 1971). United States Department of Agricul- 

ture, Animal and Plant Health Inspection Service, 

Hyattsville, Maryland. APHIS 82-3, 81 pp. 

. 1974. List of intercepted plant pests, 1972 
(pests recorded from July 1, 1971, through June 
30, 1972). United States Department of Agricul- 
ture, Animal and Plant Health Inspection Service, 
Hyattsville, Maryland. APHIS 82-4, 97 pp. 

Gressitt, J. L. 1951. Longicorn beetles of China. Lon- 
gicornia 2: 1—677. 

Hatch, M. H. 1971. The beetles of the Pacific North- 
west. Part V. Rhipiceroidea, Sternoxi, Phytophaga, 
Rhynchophora, and Lamellicornia. University of 
Washington (Seattle) Publications in Biology 16: 
1-662. 

Kobayashi, F 1985. Occurrence and control of wood- 
injuring insect damage in Japanese cedar and cy- 
press plantations. Zeitschrift fiir angewandte En- 
tomologie 99: 94-105. 

Leech, H. B. 1949. Introduction into British Columbia 
of two species of Japanese Cerambycidae (Cole- 
optera). Proceedings of the Entomological Society 
of British Columbia 45: 26. 

Maier, C. T. 1999. Beetle thought to attack only dead 


754 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


wood found in live arborvitae in Connecticut. 
Frontiers of Plant Science 51(2): 4—5. 

Makihara, H. 1984. Cerambycid beetles in Japan (5). 
Forest Pests 33: 53-54. 

Makihara, H., A. Saito, Y.-I. Chu, M. Hayashi, and S. 
Nakamura. 1989. A list of cerambycid beetles 
from Taiwan (III). Subfamily Cerambycinae. Chi- 
nese Journal of Entomology 9: 289-300. 

Mumford, B. C., compiler. 1965. List of intercepted 
plant pests, 1964 (pests recorded from July 1, 
1963, through June 30, 1964). United States De- 
partment of Agriculture, Agricultural Research 
Service, Plant Quarantine Division, Hyattsville, 
Maryland, 76 pp. 

. 1966. List of intercepted plant pests, 1965 

(pests recorded from July 1, 1964, through June 

30, 1965). United States Department of Agricul- 

ture, Agricultural Research Service, Plant Quar- 

antine Division, Hyattsville, Maryland, 88 pp. 

. 1967. List of intercepted plant pests, 1966 
(pests recorded from July 1, 1965, through June 
30, 1966). United States Department of Agricul- 
ture, Agricultural Research Service, Plant Quar- 
antine Division. ARS 82-6-1, 86 pp. 

Nakamura, S. and K. Kojoma. 1981. Immature stages 
of Taiwanese cerambycid beetles (Coleoptera: 
Cerambycidae) with notes on their habit. Kontya 
(Tokyo) 49: 155-165. 

Pu, F-J. 1991. Notes on the genus Callidiellum Linsley 
from China (Coleoptera: Cerambycidae). Acta En- 
tomologica Sinica 43: 341-343. (In Chinese.) 

Shibata, E. 1994. Population studies of Callidiellum 


rufipenne (Coleoptera: Cerambycidae) on Japa- 
nese cedar logs. Annals of the Entomological So- 
ciety of America 87: 836-841. 

Shiraki, T. 1952. Catalogue of injurious insects in Ja- 
pan. Economic and Scientific Section, Natural Re- 
sources Division. Preliminary Study No. 71 (Vol- 
ume IV). Prepared for Supreme Commander for 
the Allied Powers, Tokyo, Japan. 

USDA. 1979. List of intercepted plant pests (pests re- 
corded from July 1, 1973, through September 30, 
1977). United States Department of Agriculture, 
Animal and Plant Health Inspection Service, Hy- 
attsville, Maryland. APHIS 82-5, 568 pp. 

. 1980. List of intercepted plant pests (pests 

recorded from October 1, 1977 through September 

30, 1978). United States Department of Agricul- 

ture, Animal and Plant Health Inspection Service, 

Hyattsville, Maryland. APHIS 82-6, 178 pp. 

. 1981. List of intercepted plant pests (pests 

recorded from October 1, 1978 through September 

30, 1979). United States Department of Agricul- 

ture, Animal and Plant Health Inspection Service, 

Hyattsville, Maryland. APHIS 82-7, 171 pp. 

. 1982. List of intercepted plant pests. Fiscal 

years 1980 and 1981. United States Department 

of Agriculture, Animal and Plant Health Inspec- 

tion Service, Hyattsville, Maryland. APHIS 82-8, 

476 pp. 

. 1984. List of intercepted plant pests. Fiscal 

year 1982. United States Department of Agricul- 

ture, Animal and Plant Health Inspection Service, 

Hyattsville, Maryland. APHIS 82-9, 195 pp. 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 755-756 


NOTE 


Ectoparasitic Insects from Migrating Saw-Whet 
Owls (Aegolius acadicus) in Central Wisconsin 


A total of 833 saw-whet owls (Aegolius 
acadicus Gmelin) were trapped, providing 
an opportunity to survey these birds for ec- 
toparasitic insects. This is the largest sur- 
vey of ectoparasitic insects from this bird 
ever conducted during migration and 
should provide baseline data for future in- 
vestigators. 

Mist nets were used with a tape-recorded 
conspecific call to trap migrating saw-whet 
owls at Linwood Springs Research Station 
(44°28'’N, 89°40’W) in central Wisconsin 
during their fall and spring nocturnal mi- 
grations (IX-23 to XI-08-1997, II-18 to III- 
26-1998, IX-28 to XI-11-1998, and II-20 to 
III-27-1999). The birds were placed in tem- 
porary holding compartments for less than 
two hours. Of the 833 trapped owls, 644 
were checked for ectoparasitic insects by 
hand during banding activities. One hun- 
dred and sixty-nine of the 644 trapped owls 
were randomly selected (1/5 of the sample) 
and extensively examined for a 2-minute 
period. Collected insects were fixed in 70% 
ethanol, dehydrated through an ethanol se- 
ries to xylene, and mounted in Canada Bal- 
sam. Voucher specimens are deposited in 
the National Museum of Natural History, 
Smithsonian Institution, Washington, DC, 
accession number TM2014200. 

Sixty-four of 644 (9.9%) owls searched 
during banding and 19/169 (11.2%) when 
examined for 2 min. were positive for ec- 
toparasitic insects. Hippoboscids (costa 
americana Leach, Ornithoica vicina Walk- 
er, Ornithomyia fringillina Curtis) were har- 
bored by 1.7% of the birds, Mallophaga 
(Kurodaia acadicae Price and Beer, Strigi- 
philus acadicus Emerson and Price) by 
5.8%, and Siphonaptera (Cediopsylla sim- 


plex Baker, Orchopeas leucopus Baker) by 
3.8%. Table 1 shows the prevalence of sin- 
gle and multiple infestations during fall and 
spring migrations of owls. 

Except for hippoboscids and a single 
specimen of C. simplex (new host record), 
all ectoparasites were collected during both 
fall and spring. The three hippoboscids 
were only collected during the fall (Table 
1). As part of a larger study on Wisconsin 
hippoboscids during 11 autumns and 6 
springs, Mueller et al. (1969. Transactions 
of the Wisconsin Academy of Sciences, 
Arts and Letter 57: 189—207) collected two 
O. vicina and two O. fringillina from 234 
saw-whet owls during migration. These au- 
thors also noted that hippoboscids are un- 
common or absent during the spring. 

Orchopeas leucopus was collected from 
31 birds. Holland (1985. Memoirs of the 
Entomological Society of Canada, No. 130, 
pp. 130, 631) reported O. leucopus on 4 
saw-whet owls from Ontario, Canada. 
Specimens of this flea most likely trans- 
ferred to the owls from rodents, while our 
sole specimen of C. simplex may have 
come from a rabbit. Rodents and rabbits are 
natural hosts for these fleas. Forty-eight 
birds harbored K. acadicae and S. acadicus. 
The relatively low numbers of fleas and lice 
may be explained as follows: Fleas are in- 
termittent parasites on mammalian or avian 
hosts, and, in the case of raptors, may be 
accidental. Mallophagans complete their en- 
tire life cycle on their host but are extremely 
small, attach themselves firmly to feathers, 
and are sometimes difficult to detect on liv- 
ing birds. Additionally, lice may be present 
only as nits during bird migration (Dogiel. 
1964. In General Parasitology. Oliver and 
Boyd, Edinburgh and London 516 pp.). 


756 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
Table 1. Number of single and multiple infestations and prevalence of ectoparasitic insects from saw-whet 
owls. 
Fall Spring Fall Spring Parasite 
Ectoparasites 1997 1998 1998 1999 Totals 
Diptera 
Hippoboscidae 
Icosta americana 2/644 = 0.31% 0/644 = 0.00% 0/644 = 0.00% 0/644 = 0.00% 2 
Ornithoica vicina 6/644 = 0.93% 0/644 = 0.00% 0/644 = 0.00% 0/644 = 0.00% 6 
Ornithomyia fringillina 2/644 = 0.31% 0/644 = 0.00% 3/644 = 0.47% 0/644 = 0.00% 5 
O. vicina, O. fringillina 1/644 = 0.16% 0/644 = 0.00% 0/644 = 0.00% 0/644 = 0.00% 1 
Mallophaga 
Menoponidae 
Kurodaia acadicae 3/644 = 0.47% 2/644 = 0.31% 9/644 = 1.40% 0/644 = 0.00% 14 
Philopteridae 
Strigiphilus acadicus 10/644 = 1.55% 3/644 = 0.47% 7/644 = 1.09% 0/644 = 0.00% 20 
K. acadicae, S. acadicus 1/644 = 0.16% 1/644 = 0.16% 1/644 = 0.16% 0/644 = 0.00% 3 
Siphonaptera 
Pulicidae 
Cediopsylla simplex 0/644 = 0.00% 1/644 = 0.16% 0/644 = 0.00% 0/644 = 0.00% 1 
Ceratophyllidae 
Orchopeas leucopus 2/644 = 0.31% 8/644 = 1.24% 4/644 = 0.62% 6/644 = 0.93% 20 
Multispecies Infestations 
O. leucopus, K. acadicae 2/644 = 0.31% 2/644 = 0.31% 0/644 = 0.00% 0/644 = 0.00% 4 
O. leucopus, S. acadicus 0/644 = 0.00% 3/644 = 0.47% 0/644 = 0.00% 3/644 = 0.47% 6 
O. leucopus, K. acadicae, 1/644 = 0.16% 0/644 = 0.00% 0/644 = 0.00% 0/644 = 0.00% 1 
S. acadicus 
Seasonal Totals 30 20 24 9 83 


Stephen J. Taft, Corenna D. Kerstner, De- 
partment of Biology, University of Wiscon- 
sin-Stevens Point, Stevens Point, WI 54481, 
U.S.A. (e-mail: staft@uwsp.edu), and Eu- 


gene A. Jacobs, Linwood Springs Research 
Station, 1601 Brown Deer Lane, Stevens 
Point, Wisconsin 54481, U.S.A. 


VOLUME 102, NUMBER 3 


dvi) 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 757-758 


NOTE 


First Record of the California Pine Needle Aphid, Essigella (Essigella) 
californica (Essig) (Homoptera: Aphididae: Lachninae), in Southern Brazil 


The genus Essigella del Guercio 1909 is 
the only native Nearctic representative of 
the subtribe Eulachnina (Sorensen. 1994. 
The Pan-Pacific Entomologist 70: 1—102). 
The genus has recently been revised and in- 
cludes three subgenera, 13 species, and two 
subspecies, all of which are linear-bodied 
and feed on needles of Pinaceae, mainly Pi- 
nus, but also Pseudotsuga and Picea (Sor- 
ensen 1994; Remaudiere and Remaudiére. 
1997. Catalogue of the World’s Aphididae. 
Institut National de la Recherché Agrono- 
mique, Paris, 473 pp.). Essigella is close to 
the native Palearctic Eulachnus del Guer- 
cio, 1909, and to Pseudessigella Hille Ris 
Lambers, 1966 (Sorensen. 1990. Annals of 
the Entomological Society of America 83: 
394—408). The genus has bifid tarsal claws; 
a sclerotized dorsum; head and pronotum 
fused; meso- and metanotum fused dorsal- 
ly; abdominal tergite I usually free from the 
metanotum; and abdominal tergites HI-VII 
fused (Sorensen 1994). 

Essigella (Essigella) californica (Essig 
1909) is a lime green, small-sized (1.5—2.0 
mm) aphid found in western North America 
from southern British Columbia and Alber- 
ta to southern Mexico (Sorensen 1994; 
Blackman and Eastop. 1994. Aphids on the 
World’s Trees—An Identification and Infor- 
mation Guide. CAB International and The 
Natural History Museum, London, 987 pp. 
+ 16 pls.). One confirmed record from Mi- 
ami, Florida, suggests that it may occur in 
the Caribbean and have a pan-Mexican dis- 
tribution (Sorensen 1994). This species has 
been recently introduced into Europe. In 
France, it has been found causing damage 
on several Pinus species (Turpeau and Re- 
maudiére 1990, as cited by Sorensen 1994). 


In Spain, it has been captured by suction 
pan traps (Seco Fernandez and Mier Duarte. 
1992. Boletin de la Asociacion Espanhola 
de Entomologia 16: 255—256). 

In Brazil, E. californica was first col- 
lected by early June 1999 on slash pine Pi- 
nus elliotti Engelm. in Rolandia, Parana 
State (23°19'S, 51°22'W, altitude about 540 
m a.s.l.) and on Mexican weeping pine Pi- 
nus patula Schiede & Deppe, in the Agron- 
omy Campus of the Universidade Federal 
do Paranda, Curitiba, Parand (25°25’S, 
49°14’W, altitude 945 m as.l.) (R. C. Z. 
Carvalho, collector). Both localities repre- 
sent urban areas where the plants are used 
for landscaping. In October 1999, E. cali- 
fornica was collected at a farm in Corbélia, 
Parana (24°45’'S, 53°20'W, altitude 750 m 
a.s.l.), on P. elliotti (S. M. N. Lazzari, col- 
lector). 

Alate and apterous viviparae and nymphs 
were found on the branch tips of isolated 
plants, feeding on the base of the pine nee- 
dles, and moving quite rapidly when dis- 
turbed. Colonies of E. californica on P. pa- 
tula from Curitiba were small and associ- 
ated with Cinara pinivora (Wilson 1919) 
(Lachninae: Cinarini), while the popula- 
tions from Rolandia and Corbélia were 
more numerous, only on a few trees, and 
were not associated with other aphid spe- 
CIES: 

According to J. Sorensen (in litt.), E. cali- 
fornica is quite variable geographically, oc- 
curs on various hosts, and might be a com- 
plex. The one collected from Brazil is in the 
same phena as those which have been taken 
from Spain, France, Australia, and New Zea- 
land. It seems to be the same that occurs in 
Mendocino County, along the northern coast 


758 


of California, principally on Pinus attenuata 
Lemmon and Pinus muricata D. Don. These 
pines are in the subsection Oocarpae as is 
the Mexican weeping pine, which is closely 
related to and hybridizes well with slash 
pine in the Australis subsection. In Europe 
and Australia, E. californica occurs on Pinus 
radiata D. Don, another Oocarpae pine. In 
California, this aphid can be found on P. 
radiata, but they are less robust and much 
less common than other Essigella species. 
Sorensen also mentions that E. californica 
doesn’t do much damage, if any, on pines in 
New Zealand. 

Another Eulachnina that has been record- 
ed from Brazil feeding on Pinus, Eulachnus 
rileyi (Williams 1911) (Eastop, Costa, and 
Blackman. 1993. Pesquisa Agropecuaria 
Brasileira 28: 269-280) may be confused 
with E. californica, but they can be distin- 
guished by the following characters: E. ri- 
leyi has 6-segmented antennae, claws with- 
out bifid tips, color in life varying from 
dark olive green to gray, with a dusting of 
bluish-gray wax; E. californica has 5-seg- 
mented antennae, tarsal claws incised with 
double tips, and lime green color in life. 

Slide-mounted specimens of E. califor- 
nica are deposited in the Pe. Jesus S. Moure 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Entomological Collection, Departamento de 
Zoologia, Universidade Federal do Parana 
(DZUP). One sample of P. elliotti from Ro- 
landia is deposited under the number 8599 
and one of P. patula under the number 
8598 in the Herbarium of the Forestry De- 
partment (EFC) of the Universidade Federal 
do Parana, Brazil. 

We are thankful to Dr. John Sorensen, 
Plant Pest Diagnostics Center, California 
Department of Food & Agriculture, for con- 
firming the aphid identification and for his 
valuable comments; to Dr. David Voegtlin, 
Illinois Natural History Survey, for his 
valuable suggestions; to Dr. Jarbas Yukio 
Shimizu, EMBRAPA/Centro Nacional de 
Pesquisa de Florestas; and to Dr. Carlos 
Vellozo Roderjan, Forestry School of the 
Universidade Federal do Parana for the 
identification of the pine species. 


Regina Célia Zonta de Carvalho, Centro 
de Diagnostico Marcos Enrietti—SEAB, 
Rua Jaime Baldo 575, 80040-340, Curitiba, 
Paranda, Brasil (e-mail: regcarva@zaz. 
com.br) and Sonia Maria Noemberg Laz- 
zari, Departamento de Zoologia, Universi- 
dade Federal do Paranda, Caixa Postal 
19020, 81531-990, Curitiba, Parana, Brasil 
(e-mail: lazzari@ bio.ufpr.br). 


VOLUME 102, NUMBER 3 


759 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 759-760 


NOTE 


New Distributional and First Specific Host-Plant Records for Thionia acuta Doering 
and 7. producta Van Duzee (Auchenorrhyncha: Fulgoroidea: Issidae) 


The mostly Neotropical issid genus 
Thionia Stal (ca. 72 spp.) includes eight 
species in America north of Mexico 
(Wheeler and Wilson. 1987. Journal of the 
New York Entomological Society 95: 440— 
451). Information on host plants of North 
American Thionia species and descriptions 
of their nymphal stages have been pub- 
lished mainly in the last few years (Wheeler 
and Wilson 1987; 1988. Journal of the New 
York Entomological Society 96: 266—273; 
Wheeler. 1996. Proceedings of the Ento- 
mological Society of Washington 98: 374— 
375). Wilson et al. (1994. pp. 7-113 In 
Denno and Perfect, eds., Planthoppers: 
Their Ecology and Management, Chapman 
and Hall, New York) provided a list of re- 
corded host plants of the Fulgoroidea. Here- 
in, I provide new state records and the first 
host-plant associations for the seldom-col- 
lected T. acuta Doering, as well as the first 
specific host records for the morphological- 
ly similar T. producta Van Duzee. 


Thionia acuta Doering 


Doering (1939. University of Kansas Sci- 
ence Bulletin 25: 447—575[1938]) described 
this issid under the name 7. naso Fowler, 
relying on the identification of E. D. Ball, 
who compared Doering’s specimens from 
Concan, Tex., with Fowler’s holotype of T. 
naso from Jalapa, Mexico. Once Doering 
had access to the holotype of T. naso, she 
realized the Texas material represented a 
new species, which she named acuta (Doer- 
ing. 1941. University of Kansas Science 
Bulletin 27:185—233) and described by bib- 
liographic reference to her previous paper 
(Doering 1939). No records of 7. acuta 
have been published since Doering’s (1939) 


description, and host relationships have re- 
mained unknown. 

In 1999, I collected nymphs and adults 
of 7. acuta in Kansas and Oklahoma on ju- 
niper (Juniperus spp.; Cupressaceae) in na- 
tive stands and landscape plantings. Vouch- 
er specimens of 7. acuta (and T. producta 
from Arizona) have been deposited in the 
National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 
(USNM), and the S.W. Wilson Collection, 
Central Missouri State University, Warrens- 
burg. 

Collection records.—KANSAS: Riley 
Co., Kansas State University, Manhattan 
(39°11'39"N, 96°34'57’W; elev. 330 m), 22 
June 1999, ex Juniperus virginiana trees 
(96, 162, 3 5" instars). OKLAHOMA: 
Cleveland Co., University of Oklahoma, 
Norman (35°12’26"N, 97°26'54’W; elev. 
292 m), 12 June 1999, ex Juniperus chi- 
nensis, J. virginiana (11d, 122, 3 5™ in- 
stars); Garvin Co., unnamed road SW of 
Rts. 7 & 35, W of Davis (34°29'32’N, 
97°11'46"W; elev. 252 m), 13 June 1999, 
ex Juniperus virginiana (16, 62, 1 5 in- 
star); Woodward Co., Rt. 34, 3.7 km N of 
Vici (36°10'45"N, 99°19'55”W, elev. 690 
m), 25 June 1999 (24, 72). 


Thionia producta Van Duzee 


Described from Colorado by Van Duzee 
(1908. Proceedings of the Academy of Nat- 
ural Sciences of Philadelphia 59:467-— 
498[1907]), T. producta has since been re- 
corded from Arizona, Texas, and Utah 
(Doering 1939; Metcalf. 1958. General Cat- 
alogue of the Homoptera, Fasc. IV, Part 15 
Issidae, Waverly Press, Baltimore). The 
host plant, based on R. H. Beamer’s col- 
lecting at Leakey, Tex., on 8 July 1936, is 


760 


“cedar” (Doering 1939). My collections of 
T. producta in Arizona’s Huachuca Moun- 
tains confirm cedar (Juniperus) as a host, 
specifically alligator juniper, J. deppeana 
Steudel. 

Collection records.—ARIZONA: Co- 
chise Co., Ash Canyon Rd. nr. Twin Oaks 
Rds (0:4) iki W of (REG92 NG 123: 3IN, 
110°14.1'W; elev. 1540 m), 4 June 1997, ex 
Juniperus deppeana (96, 7¢; 1 3", 1 4", 
and 13 5" instars); Miller Canyon Rd., 1.2 
km Wor Rt 925Gil2seNe tO" l'S.2' W; 
elev. 1525 m), 5 June 1997, ex Juniperus 
deppeana (66, 12; 13 5" instars). 

Discussion.—Kansas and Oklahoma are 
new state records for T. acuta, known pre- 
viously only from Concan, Tex. The Kansas 
collection extends the range of this plant- 
hopper northeast by more than 1,100 km. 
In addition, collections of nymphs and 
adults from native stands of Juniperus vir- 
giniana and in landscape plantings from J. 
virginiana trees (10-15 m high) and a J. 
chinensis hedge (ca. 10 m long) represent 
the first published host associations for T. 
acuta. One specimen from the series that 
P.W. Oman collected at the type locality in 
Texas bears a “‘juniper”’ label, but host data 
were not mentioned by Doering (1939, 
1941). Collections of 7. producta nymphs 
from alligator juniper in the Huachuca 
Mountains of Arizona represent the first 
specific host records for a species known 
previously only from “‘cedar.”’ 

In describing 7. acuta (erroneously as T. 
naso), Doering (1939) noted its resem- 
blance to the juniper-feeding 7. producta. 
Because male genitalia of both species are 
similar—she alluded to minor differences— 
she did not describe those of T. acuta, but 
referred to the genitalic description under T. 
producta in the same publication. Although 
Doering (1939) acknowledged that some 
taxonomists might consider the two issids 
varieties of a single species, she felt it less 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


confusing if they were recognized as sepa- 
rate species. 

Doering’s (1939) hesitancy in recogniz- 
ing T. acuta and T. producta as distinct spe- 
cies might have been influenced not only 
by the morphological similarity of these 
planthoppers, but also by her awareness that 
both species had been collected on juniper 
at Concan, Tex. (assuming she saw the sin- 
gle specimen of T. acuta that bears host in- 
formation). Only in the case of 7. producta, 
however, did she publish a host association 
(Doering 1939). 

Adults of 7. acuta and T. producta, al- 
though similar, differ morphologically more 
than those of many cryptic or sibling spe- 
cies. The two species can be distinguished 
by length of the vertex (see Doering 1939: 
plate 51), even accounting for the intraspe- 
cific variation that she noted in T. producta, 
and by total body length with wings in re- 
pose (T. acuta = 6.8—8.4 mm, T. producta 
= 5.5-6.8 mm). These morphologically 
similar, juniper-feeding issids, nymphs of 
which are both green, might prove to be 
sister taxa when Thionia is analyzed cladis- 
tically. 

I thank Thomas J. Henry (Systematic En- 
tomology Laboratory, c/o National Muse- 
um of Natural History, Washington, D.C.) 
for companionship in the field, for helping 
collect issids, for comparing specimens of 
T. acuta collected in 1999 with determined 
material in the USNM, and for calling my 
attention to host information on a specimen 
of T. acuta; and Stephen W. Wilson (De- 
partment of Biology, Central Missouri State 
University, Warrensburg) for verifying (and 
in some cases correcting) my preliminary 
sorting of Thionia nymphs to instar, and for 
verifying the identification of both Thionia 
species. 


A. G. Wheeler, Jr., Department of Ento- 
mology, Clemson University, Clemson, 
South Carolina 29634, U.S.A. (e-mail: 
awhlr@clemson.edu) 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 761-767 


OBITUARY 
A TRIBUTE TO TED SPILMAN 


toe ee 
rrr rr? vy 
Ld RASH MDD 
Pr Fre + ey 
a . 


‘era. @) 7 
ow " reer Peer rer ere ry. y 
HPSS EE re rer ere es te 
Ferree CTI ress, 
a4 Pewee re rere ee 
Serireerers: tes 

eee $468 €SO845 
ereererr ee eres 
We ee ee ee ed 
j i : 
a rigeret® , 

1 +i Fi f 4 
ET ad sheds bbaade ty 
rrervree?r , G06 #4». 

Leh ded 44464008 FF" a 

' evvrreret peverryy 
444466 446 +O OP -e ss 
revvere severe rer 
shatae ae + ee! 

ger verre 


ee 
: vf 
‘ hee odes +4 
pereawererrer er ers 
2st 1444468 
Theodore James Spilman 


(1925-1996) 


762 


All who knew Theodore James Spilman 
will remember his earnest and varied inter- 
ests in many subjects, from U.S. presidents 
to music, in addition to his thorough com- 
mand of all things entomological. Ted was 
fascinated by the lives of entomologists as 
much as by entomology itself. As a devoted 
member of the Entomological Society of 
Washington, serving over the years in sev- 
eral offices including President, he had an 
unofficial role as the Society’s historian. On 
the occasion of the one-thousandth meeting 
of the ESW in February 1995, he told sto- 
ries of the early meetings and formation of 
the ESW, complete with century-old “gos- 
sip’’ and detailed knowledge of the lives of 
some founding members, leaving those of 
us present wanting to hear much more. This 
was however to be Ted’s last address to the 
membership, as an ongoing battle with can- 
cer ended his productive life in September 
1996. But Ted’s recollections and writings 
on the growth of the ESW certainly still 
give its present members, and those to 
come, a sense of their organization’s unique 
history and purpose. As fellow ESW mem- 
bers representing three generations of en- 
tomologists who knew and worked with 
Ted, we offer some personal remembrances 
of him, followed by a list of his published 
contributions to the science and history of 
entomology. 


REMINISCENCES OF TED SPILMAN 
by John M. Kingsolver 


Theodore J. Spilman died September 22, 
1996 at 71 years of age. He and I worked 
together for 23 years in our positions with 
the U.S. Department of Agriculture in 
Washington, D.C., and were the closest of 
friends. My first meeting with Ted was in 
March, 1960, while I was on a type-study 
trip to eastern museums. He and Rose Ella 
Spilman (then Warner) helped me find the 
type specimens I needed to see. By 1962, 
when I joined the U.S. Department of Ag- 
riculture staff at the National Museum, Ted 
and Rose Ella were married. The three of 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


us were part of a group of seven USDA and 
two Smithsonian Coleopterists with offices 
in the National Museum, although ento- 
mological offices and the insect collection 
were then housed temporarily in a building 
on Lamont St. north of the main museum. 
We were a congenial group although our 
respective institutional duties were some- 
what divergent. The USDA “‘crew”’ includ- 
ed Donald Anderson, George Vogt, Ted, 
Rose Ella and myself, whereas the SI group 
consisted of Oscar Cartwright and Paul 
Spangler. Later, Robert Gordon and Richard 
White joined the USDA group and Terry 
Erwin the SI group. 

Of course, I picked brains to learn pro- 
cedures, and everyone on both staffs was 
helpful. Ted and I became near “‘brothers”’ 
after we learned that he was one day older 
than I. This led to joint birthday celebra- 
tions in March for as long as we were as- 
sociated. Several years later, I discovered 
that one of my ancestors was named Spil- 
man. Although Ted tried desperately, he 
could never connect our family lines. 

Ted was well liked by everyone. He had 
a thorough knowledge of the darkling bee- 
tle family, the click beetles, and several oth- 
er smaller coleopterous families. He didn’t 
publish many large papers but seemed fated 
to put out small emergency fires that con- 
stantly cropped up in agricultural entomol- 
ogy. His papers were meticulous, practical, 
and useful. He was well-read in the arts and 
the classics, and had a considerable library 
at home. 

Ted was very professional, usually wear- 
ing a lab coat or jacket and tie whereas 
some of the others of us dressed more ca- 
sually. He was especially at home with vis- 
itors to the museum, helping them to find 
specimens in the collection, and making 
certain that they had everything they need- 
ed during their visit. 

For several reasons, Ted and Rose Ella 
and my wife and I seldom socialized. We 
had children in school, my wife worked 
evenings in a hospital, and opportunities for 
visiting were scarce. One series of social 


VOLUME 102, NUMBER 3 


gatherings thrived for several years in the 
1960’s. Several couples and singles from 
both the USDA and Smithsonian staffs 
formed a loosely organized group to meet 
and read plays. Parts would be assigned 
ahead of time, and we would meet in a 
member’s home where the plays would be 
read but not acted out. Refreshments would 
be furnished by the hosts to round out the 
evening. 

Ted often came to work at the museum 
after his retirement in 1985 although the 
last few years of his life he was seriously 
ill. I last spoke with him shortly before he 
died. 

R. J. Gagne (1997, American Entomol- 
ogist 43 (3):191—192) wrote a fine obituary 
for Ted and listed his vital statistics and 
many accomplishments, especially of his 
association with the Entomological Society 
of Washington. 


TED SPILMAN, FRIEND 
by Ross H. Arnett, Jr. 


I first met Ted when I returned to Cornell 
for a short visit soon after I went to work 
at the U.S. National Museum. V. S. L. Pate 
told me that a friend of his had a former 
student at the University of Louisville, now 
registered at Cornell, who wanted to do 
graduate work on beetle taxonomy. Pate 
wanted me to meet him. When I was in 
Comstock Hall that morning, Ted was in 
class, so I didn’t get to meet him there. I 
drove to College Town and was parking the 
car to get some lunch when this young fel- 
low ran up to the car and said: “‘You are 
Ross Arnett?” ‘Yes.’ “I am Ted Spilman 
and I want to work on Pythidae!”’ 

Thus began a close friendship that lasted 
the rest of our lives. We were in close touch 
both entomologically and personally in so 
many ways. When I left the Museum to 
teach, Ted took over my job, and the groups 
of beetles I was working on. As I wrote the 
“beetle book’’ Ted helped in so many un- 
acknowledged ways. He looked up things 
for me. He found specimens for me. He 


763 


criticized (most valuable). As everyone 
knows who has tried to do a big piece of 
work, without friends of the caliber of Ted, 
they never would complete their project. 

When I moved back to Washington to 
teach I had no place to live until we sold 
our house in Rochester. Ted put me up in 
his apartment (this was before the days of 
Rose Ella). I slept on a “‘hide-a-bed”’ in the 
living room. My head was near a fish tank. 
During the night the fish clicked their gills 
(courtship sounds?). Ted didn’t believe me. 
I don’t know if he ever found out about his 
talking fish. 

We sold our house up north. Ted and I 
went house hunting in Bladensburg. One 
day coming back from such a trip, Ted had 
an accident with his Studebaker. No one 
was hurt, but the front of the car was a 
mess. Ted never got it fixed. He drove it for 
quite awhile, and then traded it off. Ted and 
I picked out a house which my wife Mary 
and the children never saw until the day we 
moved in. 

Ted took the job in Washington before he 
finished his doctoral thesis and took his 
orals at Cornell. I suggested that he finish 
his degree at Catholic University of Amer- 
ica under my direction. He did register for 
a semester as a graduate student, but he was 
much too busy at the museum to bother 
with the trivial academic hurdles on a cam- 
pus across town, so finishing the degree 
was always put aside for something more 
important—his research. Therefore, he re- 
mained ‘‘Mr.”’ Spilman, but in our minds 
he was “Doctor.” 

Then came Rose Ella Warner. Ted and 
Rosa Ella were made for each other. It was 
my distinct honor and pleasure to be Ted’s 
best man at their wedding. Nothing between 
close friends can ever be as wonderful as 
having this role in the ceremony of the 
Holy Matrimony. I offered the toast the best 
man is supposed to do, not very elegant, I 
am afraid, but I wished them a happy mar- 
ried life, and it was to the very last. 

No matter where I moved, Ted was al- 
ways back there at the USNM (entomolog- 


764 


ical Mecca) willing to answer questions, 
send specimens, and help in every way I 
asked. He called me a few days before he 
died to say good-bye. I said “‘Good-bye, 
dear friend.” 


TED SPILMAN’S CONTRIBUTIONS TO 
ENTOMOLOGY 


by Warren E. Steiner, Jr. 


The wide range of entomological topics 
that attracted Ted Spilman’s interest, paired 
with his cordial willingness to share his 
findings with others, made him a great re- 
source to the research community at all lev- 
els. In the late 1970s when I was at the 
National Museum as a student and contract 
technician, I first got to know Ted and 
found him to be a living encyclopedia. Be- 
ing able to tap this irreplaceable source of 
information, then simply chat about topics 
of mutual interest, was then a real help to 
a beginning student, and is now truly 
missed. 

I often relied on Ted for a quick identi- 
fication, or help in finding references on a 
particular beetle or natural history topic. 
Because so many unrelated beetle taxa were 
the subjects of his research over the years, 
primarily the tenebrionoid families but also 
Ptinidae, Bostrichidae, Elateridae, Ceram- 
bycidae, and the odd little “‘jumping shore 
beetles” (intertidal Limnichidae), asking 
Ted was much faster (and more entertain- 
ing) than searching the library. 

Fieldwork was not a major part of Ted’s 
entomological career, but he did travel to 
Dominica for about 3 months in 1964 to 
participate in the Bredin-Archbold-Smith- 
sonian biological survey of the island, 
where he made some significant collections. 
Most of his collecting and work on rearing 
of larval beetles was centered in the eastern 
U.S. where he helped show me that, when 
it came to knowledge of beetles, the back 
yard was frontier. He and I made at least 
two local field trips in the early 1980s in 
search of new finds—we picked at polypore 
fungi along the Potomac, sorted through 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


sand of Delaware dunes, salvaged drowned 
specimens from the beach drift line, and 
had a good time trading trivia. But we 
turned back at Assateague Island when, to 
our dismay, we unknowingly had chosen 
the day of the annual wild pony roundup 
for our visit. Glamorous megafauna had 
overwhelmed entomology once more, but 
my adaptable travel partner was equally 
happy to find a camp along the Pocomoke, 
where the mosquitoes are at least smaller 
than those at the beach. 

Ted’s research products are a complex 
quilt of subject matter. Without hesitation 
he can be called a coleopterist. Fossil bee- 
tles, beetles of medical importance, cave 
beetles, morphology of beetle genitalia, 
beetles in mammal nests, introduction of 
foreign beetles, beetle nomenclature, rear- 
ing the larvae and pupae of beetles, the 
lives (and even grave sites) of people who 
studied beetles, were among his topics ad- 
dressed. His curatorial and identification re- 
sponsibilities covered several of the largest 
families of beetles and many unrelated 
smaller groups. He liked island faunas and 
was keeping and building separate survey 
collections of West Indian and Hawaiian 
Tenebrionidae. He seemed to enjoy solving 
complex nomenclatural problems, and in 
several papers, delved into the definition of 
terms and usage of names. The odd and 
atypical attracted his attention—he even 
named a beetle species “‘extraordinaria.”’ 

The diversity of subjects was equally 
broad when it came to the books that Ted 
reviewed. These included systematic revi- 
sions, catalogs, identification manuals, 
treatments of biology, morphology, and ag- 
ricultural importance of insects, and biog- 
raphies of entomologists. His reviews are 
still useful and amusing to read—he talks 
frankly and informally about any shortcom- 
ings and yet gives the book a good sell, 
often with humorous flair. It is obvious that 
he enjoyed reading and commenting on the 
writings of others, including works by col- 
leagues, and he sometimes published the 
same review in two or more places. 


VOLUME 102, NUMBER 3 


Ted was a long-time member of the Co- 
leopterists’ Society and served as Editor of 
the Coleopterists’ Bulletin for five years, 
succeeding Ross Arnett in 1962. With his 
writing style and acute command of taxo- 
nomic principles, terms and literature, he 
was an excellent and dependable reviewer 
of manuscripts. Alas, production of some 
taxonomic works surely must have been set 
back because of his loyal service to others 
in many ways and his constant attention to 
many duties on the job. 

A lot of Ted’s research pursuits went un- 
finished, but he leaves behind a wealth of 
information for the next students of these 
special topics, in the form of literature col- 
lections and notes, card files, manuscripts 
in progress, copies of correspondence with 
colleagues, and identified specimens. His 
sixty published works (not including book 
reviews) listed below are only the tip of a 
great iceberg of accomplishments—Ted’s 
separate files on research topics, grouped by 
subject and by taxon, will be most useful 
and unique products of his labors. These 
will perpetuate his contributions to the sci- 
ence, as will the professional example he 
set for all of us and the generations to fol- 
low. 


Publications by T. J. Spilman 


1952a. The male genitalia of the Nearctic Salpingidae. 
Coleopt. Bull. 6(1): 9-13. 

1952b. Dragonfly oviposition. The Kentucky Natural- 
ist 6(1): 15 

1953a. An odd case of gynandromorphism in the ex- 
ternal genitalia of Boros discicollis (Salpingidae). 
Coleopt. Bull. 7(5): 41—44. 

1953b. Corrections to Leng’s catalogue and Gebien’s 
Coleopterorum Catalogus: Tenebrionidae and Ka- 
talog der Tenebrioniden. J. New York Ent. Soc. 
61: 215-216. 

1954. Generic names of the Salpingidae and their type 
species (Coleoptera). J. Washington Acad. Sci. 
44(3): 85-94. 

1958. The transfer of Anamphidora from the Tene- 
brionidae to the Alleculidae (Coleoptera). Proc. 
Ent. Soc. Washington 60(6): 288. 

1959a. Notes on Edrotes, Leichenum, Palorus, Eup- 
sophulus, Adelium, and Strongylium (Tenebrioni- 
dae). Coleopt. Bull. 13: 58—64. 

1959b. A study of the Thaumastodinae, with one new 


765 


genus and two new species (Limnichidae). Co- 
leopt. Bull. 13: 111-122. 

1960a. The tenebrionid Diaclina fagi as medicine in 
the Orient. Coleopt. Bull. 14(1): 12. 

1960b. Ptinus variegatus Rossi, new to the United 
States (Coleoptera: Ptinidae). Ent. 
Washington 62(2): 103-104. 

1960c. Some synonymy in Oryzaephilus (Coleoptera: 
Cucujidae). Proc. Ent. Soc. Washington 62(4): 
Daile 

1961a. Remarks on the classification and nomenclature 
of the American tenebrionine genus Adelonia (Co- 
leoptera: Tenebrionidae). Pan-Pac. Ent. 37(1): 49— 
Syl. 

1961b. On the immature stages of the Ptilodactylidae 
(Coleoptera). Ent. News 72(5): 105-107. 

1961c. (with W. H. Anderson). On the immature stages 
of North American Pyrochroidae (Coleoptera). 
Coleopt. Bull. 15(2): 38—40. 

1961d. A few tenebrionids new to Cuba. Coleopt. Bull. 
15(4): 127. 

196le. Uloma extraordinaria, a new species from 
Cuba (Tenebrionidae). Coleopt. Bull. 15(4): 113— 
Hits). 

1962a. A few rearrangements in the Tenebrionidae, 
with a key to the genera of the Ulomini and Te- 
nebrionini of America, north of Mexico (Coleop- 
tera). Coleopt. Bull. 16: 57—63. 

1962b. The New World genus Centronopus, with new 
generic synonymy and a new species (Coleoptera: 
Tenebrionidae). Trans. Amer. Ent. Soc. 88: 1—19. 

1963a. The American genus Mycotrogus: A synopsis, 
a new species from Cuba, and a note on a larva 
(Coleoptera: Tenebrionidae). Proc. Ent. Soc. 
Washington 65(1): 21—30. 

1963b. On larvae, probably Tauroceras, from the Neo- 
tropics (Coleoptera: Tenebrionidae). Coleopt. 
Bull. 17: 58-64. 

1965. Ptinus variegatus in Georgia and Kentucky (Co- 
leoptera: Ptinidae). Proc. Ent. Soc. Washington 
67(4): 230. 

1966a. On the generic names Alphitobius, Phtora, Cla- 
moris, and Cataphronetis (Coleoptera: Tenebrion- 
idae). Proc. Ent. Soc. Washington 68(1): 6—10. 

1966b. Larva and pupa of Amarygmus morio from Ha- 
waii (Coleoptera: Tenebrionidae). Proc. Hawaiian 
Ent. Soc. 19(2): 297-301. 

1966c. A new species of Martinius from Cuba (Cole- 
optera: Limnichidae). Coleopt. Bull. 20(4): 123- 
5). 

1967a. A new North American Ulomine genus and 
species, Doliodesmus charlesi (Coleoptera: Tene- 
brionidae). Pan-Pac. Ent. 43(2): 149-154. 

1967b. Gmelin’s 13th Edition of the Systema Naturae: 
A case of neglect. Ent. News 78(7): 169-172. 

1967c. The heteromerous intertidal beetles (Coleop- 
tera: Salpingidae: Aegialitinae). Pac. Ins. 9(1): 1— 
2 


Proc. Soc. 


766 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


1968. Two new species of Niptus from North Ameri- 
can caves (Coleoptera: Ptinidae). Southw. Nat. 
13(2): 193-200. 

1969. Larva and pupa of Pyanisia tristis from Ala- 
bama (Coleoptera: Tenebrionidae). Coleopt. Bull. 
23(3): 57-61. 

1970. A new species of Tribolium from North Amer- 
ica. The results of past confusion. U.S. Dept. Agr. 
Coop. Econ. Ins. Rpt. 20(24): 396. 

1971a. Bredin-Archbold-Smithsonian Biological Sur- 
vey of Dominica: Bostrichidae, Inopeplidae, La- 
griidae, Lyctidae, Lymexylonidae, Melandryidae, 
Monommidae, Rhipiceridae, and Rhipiphoridae 
(Coleoptera). Smithsonian Contr. Zool. 70: 1—10. 

1971b. The longitudinal lines on beetle elytra: A def- 
inition of stria. Coleopt. Bull. 25(4): 121-122. 

1972a. Fossil Stichtoptychus and Cryptorama in Mex- 
ican amber (Coleoptera: Anobiidae). Univ. Calif. 
Publs. Ent. 63: 87-89. 

1972b. A new genus and species of jumping shore 
beetle from Mexico (Coleoptera: Limnichidae). 
Pan-Pac. Ent. 40(2): 108-115. 

1972c. Uloma Dejean, 1821 (Insecta: Coleoptera): Pro- 
posed designation of a type-species under the ple- 
nary powers. Z.N.(S) 1979. Bull. Zool. Nomencl. 
29(1): 32-34. 

1972d. An amendment to the proposal concerning- 
Uloma (Insecta: Coleoptera). Z.N.(S) 1979. Bull. 
Zool. Nomencl. 29(4): 194-195. 

1973a. A list of the Tenebrionidae of Michigan (Co- 
leoptera). Great Lakes Ent. 6(3): 85-91. 

1973b. Nomenclatural problems in six genera of Te- 
nebrionidae (Coleoptera). Proc. Ent. Soc. Wash- 
ington 75(1): 39—44. 

1973c. (with C. A. Triplehorn). A review of Strongy- 
lium of America north of Mexico, with descrip- 
tions of two new species (Coleoptera: Tenebrion- 
idae). Trans. Amer. Ent. Soc. 99(1): 1—27. 

1974. Temnocerus and Heterolabus, a new synonymy 
(Coleoptera: Curculionidae). Proc. Ent. Soc. 
Washington 76(3): 246. 

1976a. A new species of fossil Ptinus from fossil wood 
rat nests in California and Arizona (Coleoptera: 
Ptinidae), with a postscript on the definition of a 
fossil. Coleopt. Bull. 30(3): 239-244. 

1976b. (Witn J. D. Solomon and R. E. Doolittle). Cer- 
ambycid beetles captured in sticky-traps in Mis- 
sissippi. Coleopt. Bull. 30(3): 289-290. 

1978a. Lagria villosa in Brazil, with new descriptions 
and illustrations of the larva and pupa (Coleop- 
tera: Lagriidae). Ciencia e Cultura 30(3): 342— 
347. 

1978b. Descriptions of the larva, pupa, and adult of 
Alegoria dilatata, associated with banana plants 
in the Neotropics (Coleoptera: Tenebrionidae). 
U.S. Dept. Agr. Coop. Plant Pest Rpt. 3(6): 47-51. 

1978c. Pests not known to occur in the United States 
or of limited distribution: Lined click beetle, Agri- 


otes lineatus (L.); a wireworm, Agriotes obscurus 
(L.); a wireworm, Agriotes sputator (L.). U.S. 
Dept. Agr. Coop. Plant Pest Rpt. 3(48—52): 731-734. 

1978d. (with R. E. Woodruff and E. J. Gerberg). A false 
powderpost beetle new to the United States (Co- 
leoptera: Bostrichidae). Fla. Dept. Agr. Cons. Serv. 
Div. Plant Ind., Entomology Circular 195, 2 pp. 

1979a. (with E. J. Ford, Jr.). Biology and immature 
stages of Dirrhagofarsus lewisi, a species new to 
the United States (Coleoptera: Eucnemidae). Co- 
leopt. Bull. 33(1): 75-83. 

1979b. Larvae and pupae of Centronopus calcaratus 
and Centronopus supressus (Coleoptera: Tene- 
brionidae), with an essay on wing development in 
pupae. Proc. Ent. Soc. Washington 81(4): 513-521. 

1980. (with P. G. Crook and J. A. Novak). The lesser 
mealworm, Alphitobius diaperinus, in the scrotum 
of Rattus norvegicus, with notes on other verte- 
brate associations (Coleoptera, Tenebrionidae; Ro- 
dentia, Muridae). Coleopt. Bull. 34(4): 393-396. 

1982. False powderpost beetles of the genus Dinoderus 
in North America (Coleoptera: Bostrichidae). Co- 
leopt. Bull. 36(2): 193-196. 

1983. (with V. E Wright). An annotated bibliography 
on Prostephanus truncatus (Horn), (Coleoptera: 
Bostrichidae): A pest of stored grain. Trop. Stored 
Prod. Inf. 46: 25—30. 

1984a. Vignettes of 100 years of the Entomological 
Society of Washington. Proc. Ent. Soc. Washing- 
ton 86(1): 1-10. 

1984b. Identification of larvae and pupae of the larger 
grain borer, Prostephanus truncatus (Coleoptera, 
Bostrichidae), and the larger black flour beetle, 
Cynaeus angustus (Coleoptera: Tenebrionidae). 
Proc. Third Intl. Working Conf. Stored-Product 
Ent. 44-53. 

1985. Review of Lanelater of North America (Cole- 
optera: Elateridae). Insecta Mundi 1(1):1—12. 
1989a. A visit to the tomb of Thomas Say in New 
Harmony, Indiana. Ent. News 100(5): 224—227. 

1989b. Vignettes of the Presidents of the Entomolog- 
ical Society of America, 1889-1989. Bull. Ent. 
Soc. Amer. 35(3): 33-65. 

1991a. (with J. E Lawrence). Tenebrionidae (Tene- 
brionoidea) (including Alleculidae, Cossyphodi- 
dae, Lagriidae, Nilionidae, Rhysopaussidae, Ten- 
tyriidae), pp. 520-528. Jn Stehr, E W., ed., Im- 
mature Insects. Kendall-Hunt, Dubuque, Iowa. 

1991b. (In Gorham, J. R., Insect and mite pests in 
food) Part 1.6. Spider beetles (Ptinidae, Coleop- 
tera). U.S. Dept. Agr. Handbook 655(1): 137-147. 

1991c. (In Gorham, J. R., Insect and mite pests in 
food) Part 1.11. Darkling beetles (Tenebrionidae, 
Coleoptera). U.S. Dept. Agr. Handbook 655(1): 
185-214. 

1995. (with R. L. Aalbu and K. W. Brown). The sys- 
tematic status of Amblycyphrus asperatus, Thren- 
us niger, Pycnomorpha californica, Emmenastus 


VOLUME 102, NUMBER 3 


rugosus, and Biomorphus tuberculatus Motschul- 
sky (Coleoptera: Tenebrionidae). Proc. Ent. Soc. 
Washington 97(3): 481—488. 


ACKNOWLEDGMENTS 


For discussions and comments on Ted’s 
“life history” and reviews of early drafts 
of this remembrance, we thank D. M. An- 
derson, R. J. Gagné, Stuart E. Neff, Paul 
and Phyllis Spangler. 


767 


Warren E. Steiner, Jr., John M. Kingsol- 
ver, and Ross H. Arnett, Jr. (WES) Depart- 
ment of Entomology, NHB-187, Smithsoni- 
an Institution, Washington, DC 20560, 
U.S.A.; (JMK) Florida State Collection of 
Arthropods, P.O. Box 147100, Gainesville, 
FL 32614, U.S.A.; (RHA), 2406 NW 47th 
Terrace, Gainesville, FL 32606, U.S.A. (de- 
ceased). 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 768-770 


Book REVIEW 


The Genus Rhipicephalus (Acari, [xodi- 
dae): A Guide to the Brown Ticks of the 
World. Jane B. Walker, James E. Keirans 
and Ivan G. Horak. Cambridge Univer- 
sity Press, The Edinburgh Building, 
Cambridge CB2 2RU, United Kingdom. 
Kit (643 4pp..7.27 9ehies/ de tabs:, 74 
maps. 2000. $100.00/cloth. ISBN 0-521- 
48008-6. www.cup.cam.ac.uk. 


Through nearly two centuries of study, 
no tick genus has proved as taxonomically 
intractable as Rhipicephalus. Particularly in 
tropical Africa, exclusive home to 60 of the 
world’s 74 known species, misidentifica- 
tions of rhipicephalids have rendered sus- 
pect numerous otherwise valuable veteri- 
nary and parasitological papers. Now, how- 
ever, as if in tribute to the new millennium, 
acarology’s own Gordian knot lies sundered 
by the peerless team of Drs. Walker (On- 
derstepoort Veterinary Institute, South Af- 
rica), Keirans (U.S. National Tick Collec- 
tion, Georgia Southern University, States- 
boro) and Horak (Faculty of Veterinary Sci- 
ence, University of Pretoria, South Africa), 
whose decades-long synergy has yielded 
the most comprehensive—and attractive— 
volume ever published on a single genus of 
ticks, the sixth in a series of monographs 
on the Ixodoidea begun by George Henry 
Falkiner Nuttall (1862-1937) and col- 
leagues in 1908 and championed from the 
outset by Cambridge University Press. 

Introductory sections of this sumptuous 
work provide a definition of Rhipicephalus 
in the context of the family Ixodidae (sec- 
tion 2), augmented by illustrations of key 
characters (section 3) and a glossary (sec- 
tion 4) of all morphological, taxonomic and 
other terms that are associated with mem- 
bers of this genus (some of these, like “‘sha- 
greened” (p. 19), are uniquely rhipicephalid 
descriptors). There follows (section 5) a 
worldwide annotated checklist of all the 
Rhipicephalus species names ever pub- 


lished, including junior synonyms, new 
combinations and lapsus calamorum, to- 
gether with full citations for original de- 
scriptions, and collection data and deposi- 
tories for types and type series. The book 
then neatly divides itself in two: sections 6, 
7 and 8 describe the rich rhipicephalid fau- 
na of the Afrotropical Zoogeographic Re- 
gion; sections 9, 10 and 11 address the 10 
Rhipicephalus species that occur outside the 
Afrotropics. As noted by the authors, this 
dichotomy does not work for all taxa. In- 
cluded in their Afrotropical discussion is 
the circumglobal R. sanguineus, the world’s 
most widespread tick species and the only 
representative of its genus in the Western 
Hemisphere, as well as R. fulvus, most col- 
lections of which have come from Tunisia 
(i.e., the Palearctic Zoogeographic Region), 
and R. camicasi and R. turanicus, with vast 
ranges in Africa and adjacent lands. Also, 
the trans-African subspecies R. e. evertsi 
apparently has been recently introduced by 
humans into Yemen and the Red Sea coast 
of Saudi Arabia. But for the most part, the 
Rhipicephalus faunas of Africa and the rest 
of the world do not intersect, and the pre- 
liminary discussions of each (sections 6 and 
9) comprise fascinating historical reviews 
of regional tick research, exhaustive lists of 
country-specific references, and the au- 
thors’ long-anticipated male and female 
identification keys, whose user-friendliness 
is immeasurably enhanced by inclusion of 
information on preferred hosts and distri- 
butions within the key couplets themselves. 

At the core of this effort are the individ- 
ual species accounts (sections 7 and 10), 
each wholly self-contained, which collec- 
tively constitute over three quarters of the 
text. In these accounts, thorough descriptive 
diagnoses of all known active stages are ac- 
companied by stunning scanning electron 
photomicrographs, conventional light mi- 
crographs of slide-mounted female genital 


VOLUME 102, NUMBER 3 


apertures a la Feldman-Muhsam (1956, 
Bull. Res. Council Israel 5B: 300—306), 
and, for both sexes, the heartbreakingly 
beautiful pen-and-ink drawings of A. Ol- 
wage, Walker’s long-time artistic collabo- 
rator. Olwage’s preeminence as a tick illus- 
trator gained world renown with the publi- 
cation of his series of color plates depicting 
the adults of Amblyomma species capable 
of transmitting the agent of heartwater, 
Cowdria ruminantium (1987, Onderste- 
poort J. Vet. Res. 54: 353-379). In the opus 
at hand, his skill is perhaps best exemplified 
in drawings of the ornate R. pulchellus, 
whose specific epithet means “beautiful” 
(p. 366), and the densely punctate male of 
R. ramachandrai (p. 562). Olwage also pro- 
duced the final versions of each tick distri- 
bution map, wherein data as diverse as type 
locality Gf known), and confirmed and un- 
confirmed records of various authors are 
precisely but harmoniously plotted (excel- 
lent examples are the distribution of R. guil- 
honi, p. 207, and R. sulcatus, p. 431). Given 
the biomedical importance of this genus, it 
is surprising how many diagnoses are in- 
complete for at least one life history stage. 
While males and females are known for all 
species, both nymphs and larvae are un- 
known for the Afrotropical R. aquatilis, R. 
bequaerti, R. bergeoni, R. boueti, R. deltoi- 
deus, R. dux, R. interventus, R. longiceps, 
R. masseyi, R. oreotragi (one of two new 
species described in this book; the other is 
R. warburtoni, known from all stages), R. 
punctatus, R. serranoi and R. supertritus. In 
addition, the nymph of R. longicoxatus is 
unknown, as is the larva of R. pseudolon- 
gus. Preimagines of R. complanatus and R. 
ziemanni exist in collections but have not 
been described. Outside Africa, the imma- 
tures of R. scalpturatus are unknown, as 
well as the larva of R. leporis, while the 
larva of R. pusillus was not available for 
study. 

Each species diagnosis closes with one or 
more paragraphs of critically important 
“notes on identification,” in which the au- 
thors offer insights for separating similar 


769 


species, often discerning helpful differences 
in minor morphological expressions, such 
as those accompanying engorgement, or in 
tick ecology, distribution, and host specific- 
ity. This discussion leads seamlessly to a 
tabulation of all known hosts of each tick 
species, these data having been gleaned 
from large and small tick collections world- 
wide and from the literature (hosts are en- 
tirely unknown for R. deltoideus and R. 
scalpturatus). Statements on tick zoogeog- 
raphy, incorporating altitudinal data, rain- 
fall and vegetation zones, and often ex- 
plaining or clarifying the map records, 
round out each account, together with a 
summary of disease relationships (only 27 
Rhipicephalus species (36%) have ever 
been linked to any human or animal dis- 
ease, several transmitting only in the labo- 
ratory to splenectomized hosts) and a list of 
species-specific references. At the close of 
both the African and non-African species 
accounts, the tick/host tables of the ac- 
counts are combined and inverted, yielding 
checklists of all Rhipicephalus species col- 
lected from each host animal (sections 8 
and 11). In this reversed arrangement, tick 
names in boldface generally indicate that 
the animal under which they are listed is a 
preferred host of the adults. However, in the 
Afrotropical R. follis, R. gertrudae and R. 
simus, the hosts of immatures (usually 
small rodents) are entirely unrelated to the 
hosts of adults (usually ungulates), so these 
three tick species appear twice at opposite 
ends of the checklist. With additional field 
collections of immatures, this bimodal pat- 
tern of parasitism is likely to apply to many 
more rhipicephalids. Dual entries also ap- 
pear for the immatures of the Afrotropical 
R. pravus, R. sp. near pravus and R. war- 
burtoni, which are often found on lago- 
morphs (like most African Rhipicephalus) 
but chiefly parasitize elephant shrews. 
Section 12, ““Species groups based on the 
immature stages,”’ is a first attempt to help 
taxonomists identify the maddeningly sim- 
ilar immatures of this genus by sorting 
known nymphs and larvae into 10 more or 


770 


less recognizable groups (8 Afrotropical, 2 
non-Afrotropical) based on diagnostic ca- 
pitular characters, again admirably drawn 
by Olwage. Brief descriptions of these char- 
acters are provided for each group and, as 
in the adult key, summaries of each species’ 
geographic distribution assist in clinching 
identifications. The authors take pains to 
note that these groups may not be “natu- 
ral,’ although the immatures of the Afro- 
tropical R. evertsi group—R. bursa, R. ev- 
ertsi and R. glabroscutatum—are both mor- 
phologically similar and the only species in 
their genus known to have a two-host, rath- 
er than three-host, life cycle. As well, spe- 
cies in the ‘“‘pointy-palp”’ groups of nymphs 
and larvae (1.e., the Afrotropical R. capen- 
sis, R. follis and R. simus groups), which 
are exceedingly difficult to identify, seem 
to share the habit of feeding on specific 
small mammals that are not parasitized by 
the adult stages. 

To describe in any detail the variety of 
diseases associated with members of the ge- 
nus Rhipicephalus, and the research that has 
been conducted in this sphere, would ne- 
cessitate doubling the present volume. This 
is, after all, a book about ticks: how to iden- 
tify them, how they live, and where and 
when they may be found. But just as the 
authors employ host/tick checklists to sum- 
marize the discussions of host relationships 
in the individual species accounts, in their 
final section (13) they leave disease-orient- 
ed readers with two multipage, landscape- 
format tables on the transmission of diseas- 
es to animals and humans. Both are superb 
synopses. The animal table lists 25 Rhipi- 
cephalus-borne diseases and 28 causative 
agents, plus two afflictions (tick paralysis 
and tick toxicosis), with additional headers 
for the animals affected, the tick vectors 
(including, if known, the primary natural 
vectors and any that have transmitted ex- 
perimentally), the number of hosts utilized 
by each tick species (in Rhipicephalus gen- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


erally 3, sometimes 2), the tick stages that 
acquire and transmit disease, and one or 
more classic references (an appended bib- 
liography provides 80 additional reviews of 
animal diseases vectored by rhipicephalid 
ticks). The table for tickborne diseases of 
humans is similar, listing 15 diseases and 
causative agents, with further headers for 
tick vectors, number of hosts, the mode of 
infection or isolation of the causative 
agents, and classic references (the appended 
bibliography for this table provides 33 ad- 
ditional reviews). 

Even at the dawn of the 21st century, 
books of this length and complexity are cer- 
tain to contain at least a few minor typo- 
graphical errors. Accordingly, the publish- 
ers have agreed to issue an erratum slip, 
which currently makes the following emen- 
dations: the correct date of publication for 
the new species R. oreotragi (p. 31) and R. 
warburtoni (p. 38) is 2000, the date 1999 
being an unfortunate carryover from pub- 
lication delays; the Rocky Mountain Lab- 
oratory is in Montana, a state that should 
be abbreviated MT, not MA (Massachu- 
setts) (p. 45); and under the “notes on iden- 
tification’”’ for R. serranoi (p. 408), “XX” 
and ‘“‘YY”’ should be replaced by male and 
female symbols. Such minutiae aside, this 
authoritative and beautiful book is the best 
beginning we could wish for acarology’s 
third century. No one who visits its pages 
will fail to find instruction or inspiration. 
Surely somewhere Cecil Warburton (1854— 
1958), whose meticulous studies of Rhipi- 
cephalus intraspecific variation made him 
the obvious choice for this work’s dedica- 
tory frontispiece, is smiling serenely at his 
reward: an infinite number of ticks and an 
eternity in which to examine them. 


Richard G. Robbins, Armed Forces Pest 
Management Board, Walter Reed Army 
Medical Center, Washington, DC 20307- 
5001, U.S.A. (E-mail: robbinrg @acq.osd. 
mil) 


PROC. ENTOMOL. SOC. WASH. 
102(3), 2000, pp. 771-772 


INSTRUCTIONS 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- 
MMEMt CO ay ons 8 (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. 


NIA NS 


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 < 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 in the text. Reference in text 
shouldiberas = “Fig. I’ 2 Bicss 293) .a- a= 
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, 
Ele: 

When referring to references in text, use 
the following; Henry (1990), (Henry 1990), 
(Henry 1987, 1990), (Smith 1990, Henry 
1992), (Smith 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 


UY 


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 


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 


Grissell and Michael E. Schauff. 87 pp. 1997 


MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Memoirs 2, 3, 7, 9, 10, 11, and 13 are no longer available. 


No. 
No. 
No. 
No. 


No. 


No. 


No. 


No. 


No. 
No. 


No. 


he: 


4. 
5: 
6 


nl. 


22: 


The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 
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 
i eeetee ANE LENO tac deer oak 0 BN. Tes AO epee ee ate cee ms ae aa 


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 _ 


. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. 


PST UEC, CEPR Gis OSES ah GSU ERC LRA G i SONS LCE SAP ee aI ah Raed Pee AU ae 5 AIR 


. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174 


SS poh SSE a Sg Td eh rad Ry ME SBR ee Sa of SURES. 1) EOD EM Ey Bl RN ER 


. A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent 


Benes ar padee 1 LOS pp POG ai sai ew Mien LE yo ied Se i ty als 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 


ee Orden 2ndvAL SAMvienks” 26 1p. POOG sre oe 2m fae a EES 


. Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and 


emitter Oe Wey PP TOR ox eee Oy 8E) ie a eI Ao ale we saan A) 


. Monograph of the Stilt Bugs, or Berytidae (Heteroptera), of the Western Hemisphere, by 


STEP EIS bd Sen) ets 12 Ji ea a ae TAR pes oda a Meg, stake SA OE ae SOE. AT A eee eee Oye ea OS 


. The Genera of Elaphidiini Thomson 1864 (Coleoptera: Cerambycidae), by Steven W. Lin- 


SEE USE plo aye TEE 2 Cog RONEN Uo cbs Ule VAS ab ea tn wots Spree MERLE Cs CR Ame MERE PP alco, YA AS me 


New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae), by David G. 
(Bie AUS oe GS Pan NON oe aie A SBb ieee Napa br Ole Ne OR RUD oS Re ON Ela OER Me DO athe” Peer 0. la Dae 


Systematics of the North American Species of Trichogramma Westwood (Hymenoptera: 
Pnucvoecranimatidas). by John. Pinte, 28 7pp, 1999 2 Na 


$10.00 


15.00 


$15.00 
15.00 
15.00 


15.00 


12.00 


5.00 


25.00 


25.00 
25.00 


25.00 


25.00 


18.00 


12.00 


12.00 


28.00 


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 
percent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the 
Custodian, Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 
Washington, D.C. 20560-0168. 


CONTENTS 


(Continued from front cover) 


MATHIS, WAYNE N. and JAMES F. EDMISTON—A revision of the shore-fly genus Lemnaphila 
Cresson (Diptera: Ephydridae) 


PAPP, JENO and SCOTT R. SHAW—A study of the genus Falcosyntretus Tobias from the 
New World with five new species and a key to known species (Hymenoptera: Braconidae: 
Euphorinae) 

PIKE, K. S., P. STARY, T. MILLER, G. GRAF, D. ALLISON, L. BOYDSTON, and R. MILLER— 
Aphid parasitoids (Hymenoptera: Braconidae: Aphidiinae) of northwest USA 


POLAVARAPU, S., JOHN A. DAVIDSON, and DOUGLASS R. MILLER—Life history of the 
Putnam scale, Diaspidiotus ancylus (Putnam) (Hemiptera: Coccoidea: Diaspididae) on blue- 
berries (Vaccinium corymbosum, Ericaceae) in New Jersey, with a world list of scale insects 
on blueberries 


SITES, ROBERT W. and MICHAEL R. WILLIG—Morphometric variation among populations of 
Ambrysus mormon Montandon (Heteroptera: Naucoridae) 


VASU, V., DAVID R. SMITH, and MALKIAT S. SAINI—Review of the Asian sawfly genus 
Anisoarthra Cameron (Hymenoptera: Tenthredinidae) 


NOTES 


TAFT, STEPHEN J., CORENNA D. KERSTNER, and EUGENE A. JACOBS—Ectoparasitic 
insects from migrating saw-whet owls (Aegolius acadicus) in central Wisconsin 


WHEELER, A. G., JR.—New distributional and first specific host-plant records for Thionia acuta 
Doering and T producta Van Duzee (Auchenorrhyncha: Fulgoroidea: Issidae) 


ZONTA DE CARVALHO, REGINA CELIA, and SONIA MARIA NOEMBERG LAZZARI— 


First record of the California pine needle aphid, Essigella (Essigella) californica (Essig) 
(Homoptera: Aphididae: Lachninae), in southern Brazil 


OBITUARY § 


STEINER, WARREN E., JR., JOHN M. KINGSOLVER, and ROSS H. ARNETT, JR.—A tribute 
to Ted Spilman 


BOOK REVIEW 


ROBBINS, RICHARD G.—The Genus Rhipicephalus (Acari, Ixodidae): A Guide to the Brown 
Ticks of the World, by Jane B. Walker, James E. Keirans, and Ivan G. Horak 


MISCELLANEOUS 


Instructions for authors 


549 


533 


601 


| VOL. 102 OCTOBER 2000 NO. 4 
: QL. (ISSN 0013-8797) 


HG | 


© PROCEEDINGS 


» 
4 


| of the 


ENTOMOLOGICAL SOCIETY 


CONTENTS 


ALUJA, MARTIN, JAIME PINERO, MAURILIO LOPEZ, CESAR RUIZ, ALBERTO ZUNIGA, 
ENRIQUE PIEDRA, FRANCISCO DiAZ-FLEISCHER, and JOHN SIVINSKI—New host 
plant and distribution records in Mexico for Anastrepha spp., Toxotrypana curvicauda 
Gerstacker, Rhagoletis zoqui Bush, Rhagoletis sp., and Hexachaeta sp. (Diptera: Tephritidae) 802 


BORKENT, ART—The larva and pupa of Schizonyxhelea forattinii Wirth and Grogan (Diptera: 
Ceratopogonidae) with a discussion of the phylogenetic relationships of the genus .......... 862 


BRANHAM, MARC A. and MIGUEL ARCHANGELSK Y—Description of the last larval instar 
and pupa of Lucidota atra (G. A. Olivier 1790) (Coleoptera: Lampyridae), with a discussion 
Gidodomuinal sepment homology across life stages) )) 22.2020. dices we enna voce eee edesedsevse 869 


BROWN, JOHN W. and JON LEWIS—Catalogue of the type specimens of Tortricidae (Lepidop- 
tera) in the collection of the National Museum of Natural History, Smithsonian Institution, 


WTSS HR EEAWOPeS ID OMS Se GS a ea As eee Se eRe a DL SAGY 4 eR apa RTRSY MR ed AP Ue ete A at 1014 
CONTRERAS-RAMOS, ATILANO—A new species of Chloronia Banks (Megaloptera: Corydal- 

idae) from southeastern Brazil, with a key to the species of Brazil .......................4.. 919 
DARSIE, RICHARD FE, JR.—Description of the pupa of Armigeres (Leicesteria) omissus (Edwards) 

and a key to the larvae and pupae of the Armigeres occurring in Nepal (Diptera: Culicidae) .. 964 
ECKERLIN, RALPH P. and HARRY F. PAINTER—New records of fleas (Siphonaptera) from 

SSISURSIND) VIWEG NATE SCE Ne Sg ea Re yn oe aNd A ea Papa Neer WES Aha ae 969 


EVANS, HOWARD E.—Three new species of Dipogon Fox (subgenus Dipogon) (Hymenoptera: 
Pompilidae) from central and western North America ................ 0.02 cece eee cece eee eens 1010 


GAGNE, RAYMOND J., HELGA BLANCO-METZLER, and JEAN ETIENNE—A new 
Neotropical species of Clinodiplosis (Diptera: Cecidomyiidae), an important new pest of culti- 
Seca pels (Capsicuin Spps:, SOlAnACEAG), (02 pahie slo ekret Nafatac se cies eae ate emclayh aie Sine 4 abscissa 831 


GOEDEN, RICHARD D.—Life history and description of immature stages of Neaspilota pubescens 
Freidberg and Mathis (Diptera: Tephritidae) on Lessingia filaginifolia (Hooker and Arnott) M. A. 


Mave i AStErACcAC) Min SOULAe Ti CallfOrnlay., oleic :.°4 2 Sake ayes tials Sisiele Beis Ret sete ek ae Ge Gisele es ce seicet 878 
GUILBERT, ERIC—Revision of the genus Parada Horvath (Hemiptera: Tingidae) with cladistic 
SLT STIS 92 sai hk Tah os Baad Se NRG to ANS Boa ORI AT (5 gg a ae 816 


(Continued on back cover) 


THE 


ENTOMOLOGICAL SOCIETY 
OF WASHINGTON 


OFFICERS FOR 2000 ; 
Davip G. Furtu, President MICHAEL G. POGUE, Treasurer 
JoHN W. Brown, President-Elect THEODORE R. Scuuttz, Program Chair 
Stuart H. McKamey, Recording Secretary STEVEN W. LINGAFELTER, Membership Chair 


Ho us B. WILLIAMS, Corresponding Secretary MicuHaet E. Scuaurr, Past President 
Jon A. Lewis, Custodian 


Davip R. Smitn, Editor 


Publications Committee 


Ss ee ee 


RAYMOND J. GAGNE THOMAS J. HENRY Wayne N. MaArTHIis 
Honorary President 
Louise M. RussELL 
Honorary Members 
KARL V. KROMBEIN RONALD W. HopGEs 
DoNALD M. ANDERSON WILLIAM E. BICKLEY 


All correspondence concerning Society business should be mailed to the appropriate officer at the following 
address: Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Wash- 
ington, D.C. 20560-0168. > 
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 7:30 P.M. Minutes of meetings — 


ce ae 


are published regularly in the Proceedings. ti 
MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology. — 
Annual dues for members are $25.00 (U.S. currency). 's 


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, Smithsonian H 
Institution, Washington, D.C. 20560-0168. Members in good standing receive the Proceedings of the Entomo- 
logical 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. i 
The Society does not exchange its publications for those of other societies. 4 
PLEASE SEE PP. 771-772 OF THE JULY 2000 ISSUE FOR INFORMATION REGARDING | 
PREPARATION OF MANUSCRIPTS. a 
STATEMENT OF OWNERSHIP i 

Title of Publication: Proceedings of the Entomological Society of Washington. Le 
Frequency of Issue: Quarterly (January, April, July, October). i ; 


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 4 
ington, D.C. 20560-0168. 

Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomologysg® 
Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. 

Books for Review: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of — 
Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. 


Managing Editor and Known Bondholders or other Security Holders: none. 


putes 


= 


io On SS, BE 


This issue was mailed 17 October 2000 
Periodicals Postage Paid at Washington, D.C. and additional mailing office. 


ay 


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


This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). 


t 


Se eee enor 


PROC. ENTOMOL. ; 
102(4), 2000, ; 


“”. WASH. 
773-193 


FLOWER FLIES OF THE SUBGENUS OCYPTAMUS (MIMOCALLA HUi- 
(DIPTERA: SYRPHIDAE) 


FE CHRISTIAN THOMPSON AND MANUEL A. ZUMBADO 


(FCT) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. 
Department of Agriculture, % National Museum of Natural History Smithsonian Insti- 
tution, Washington, DC 20560-0169 U.S.A. (e-mail: cthompso@sel.barc.usda.gov); 
(MAZ) Entomology, Instituto Nacional de Biodiversidad, Santo Domingo, Heredia, Costa 
Rica (e-mail: mzumbado @inbio.ac.cr) 


Abstract.—The subgenus Ocyptamus (Mimocalla) is revised. Ocyptamus (Mimocalla) 
tristani Zumbado, new species, is described and two new synonyms are proposed (Sal- 
pingogaster flukei Curran 1941 and Baccha polista Hull 1943 = Baccha bonariensis 
Bréthes 1905). A key to the species, along with synonymies, descriptions, distributions, 


and figures for all species are given. 


Key Words: Neotropical, key 


The New World tropics have the richest 
diversity of flower flies in the World. This 
diversity holds great promise for sustain- 
able agriculture. The flower fly diversity 
largely resides in three genera (Toxomerus 
(150 spp.), Ocyptamus (300 spp.) and Co- 
pestylum (320 spp.)). The flies of the genus 
Ocyptamus are predators of a diverse array 
of plant pests (mites, scales, plant hoppers, 
aphids, etc.). To fully utilize their potential, 
the species must be named and described 
so that their life-histories can be studied and 
reported. Then the relative contribution of 
these predators to agroecosystems can be 
assessed and, perhaps, enhanced. 

Consider coffee production: While most 
coffee is today grown in large, open, single 
crop systems which require insecticide use, 
there is a growing trend to return to more 
natural shade-grown systems. In such sys- 
tems, the new species described herein may 
be a useful biological control agent. The 
species is a predator of black, soft-brown 
and hemispherical scales, which are pests 
of coffee and various ornamental plants. 


This paper is the second in a series to 
treat various components of the large di- 
verse genus Ocyptamus. The first part 
(Thompson, in preparation) includes a de- 
scription and diagnosis of the genus and a 
key to the components (subgenera and spe- 
cies groups) along with the treatment of 
several species groups. This paper treats the 
species group named and treated as a sub- 
genus by Hull. Until a cladistic analysis is 
done of the genus as a whole, we, for prag- 
matic reasons, retain Mimocalla as a sub- 
genus. The likely sister to Mimocalla is O. 
conjunctus Wiedemann and the sister to O. 
conjunctus + Mimocalla is O. sargoides 
Macquart. Using these species as out- 
groups, we include a preliminary cladistic 
hypothesis for the species of Mimocalla. 
Redescriptions, complete synonymies, illus- 
trations, and distributions are given for all 
species. Terminology follows Thompson 
(1999), the abbreviations found in the syn- 
onymies follow Thompson (2000), the use 
of the asterisk in the distribution statement 
means verified records to be found below 


774 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Habitus of Ocyptamus tristani, dorsal. 


in the material examined section. Color im- 
ages of these flies along with the rest of the 
information in this paper may be found on 
the Diptera WWW site (http://www.sel. 
barc.usda.gov/Diptera/) and on the Diptera 
Data Dissemination Disk. 


Genus Ocyptamus 
Ocyptamus Macquart 1834: 554. Type spe- 
cies, fascipennis Macquart (Coquillett 
1910: 577) = fuscipennis Say. 
Subgenus Mimocalla Hull 


Baccha subg. Mimocalla Hull 1943d: 46. 
Type species, Baccha capitata Loew 


T. Litwak 


(orig. des.); Hull 1949: 97 (discussion, 
key). 


Description.—Face usually yellow, rare- 
ly with weak brownish vitta or black with 
narrow yellow vitta, with strong tubercle; 
frons prominent; vertex short, broad, with 
ocellar triangle about 1% times its length 
anterior to hind margin of eye; occiput with 
2 rows of cilia, with cilia normal, not scale- 
like; metasternum bare; metathoracic epis- 
ternum pilose; postmetacoxal bridge incom- 
plete; scutellum yellow or rarely entirely 
black, with 1 row of strong and long ventral 


VOLUME 102, NUMBER 4 


pile; plumula long to absent; calypter with 
strong fringe only on ventral edge; meson- 
otum usually with bright yellow pollinose 
vittae, rarely entirely dark; metatrochanter 
always with a tuft of long black pile, fre- 
quently males with long ventral pile also on 
metacoxa and femur; abdomen clavate, 
constricted at base of 2nd tergum and grad- 
ually widening apically, brown to black, 
with yellow fasciae; male genitalia greatly 
enlarged, with cercus and apicolateral cor- 
ners of 9th tergum enlarged; male 5th ster- 
num variously modified on apical half. 

Wing: Alula normal, about 1% times as 
broad as cell BM, bare or microtrichose; 
crossvein r-m at basal % of cell DM, slight- 
ly beyond termination of vein Sc; vein M1 
sinuate, ending approximately at termina- 
tion of vein R1; vein R4+5 sinuate, making 
a shallow loop into celi R4+5. 

Length: 12-16 mm; wing, 10—14 mm. 

Discussion.—The subgenus Mimocalla is 
readily distinguished from all other groups 
of Ocyptamus by: 1) strongly petiolate and 
clavate abdomen; 2) large size; 3) enlarged 
male genitalia including a modified 4th 
sternum; 5) vein R4+5 sinuate; 6) vein M1 
sigmoid; and 7) metatrochanter with a 
strong pile tuft. In appearance Mimocalla 
can only be confused with O. (Theranta) 
conjunctus, but Mimocalla is readily distin- 
guished by the sinuate vein R4+5 and short 
antenna. 

While a full cladistic analysis has not 
been done, the characters of the male gen- 
italia suggest these relationships (Mimocal- 
la = (giganteus + willistoni) + ((bonarien- 
sis + erebus) + (nymphaea + (capitatus + 
tristani)))). In most Ocyptamus, the cercus 
is unmodified, small, and oval, but in Mim- 
ocalla the cercus is enlarged. In one group 
(giganteus + willistoni), the shape of the 
cercus is triangular and the surstylus short, 
blunt apically and directed ventrally. In the 
other group, the cercus is elongate and the 
surstylus is also elongate, tapered apically 
and directed posteriorly. Within this second 
group, the genitalia of capitatus and tristani 
are the same, and those of nymphaea are 


ads 


very similar. These three species als. hare 
the more or less simple 5th sternum. “he 
other two species (bonariensis, erebis) 
have the 5th sternum produced apicolater- 
ally on the right side. 

Little is known about the biology of 
Ocyptamus species. However, the life his- 
tory of one species of Mimocalla has been 
published. We review that work in light of 
our rearing of tristani, new species. Bruch 
(1923) noted that bonariensis larvae com- 
pletely suck dry a scale insect before mov- 
ing on to the next and that the complete 
larval cycle probably lasts about three 
weeks. Once the larva finishes feeding, the 
larva remains quiescent for about two days, 
then evacuates the gut and changes position 
in order to pupate, either on the same plant 
or nearby, and finally resting about one 
more day before pupating. The larva slowly 
becomes smaller and releases a viscous 
transparent liquid, which when hardens to 
anchor the puparium. During the process of 
transformation the posterior part of the lar- 
va becomes narrower, while undulatory and 
contraction movements of the pupa forming 
inside inflates bit by bit the anterior portion; 
this process lasts about two hours. The re- 
cently formed puparium is colored the same 
as the mature larva but shortly after pupa- 
tion it becomes pale brown. The pupal stage 
lasts about two weeks. 

Observations by M. Zumbado and J. A. 
Ugalde on tristani confirm Bruch’s obser- 
vations. Mature tristani larvae (Figs. 20-— 
22) measure between 12 and 14 mm long 
and about 5 mm wide. The body is subpar- 
allel along its length, rather convex dorsally 
and flat ventrally; anteriorly it is narrower 
and posteriorly is obliquely truncated. The 
color is similar to that of bonariensis. The 
puparium (Fig. 23) is smaller than the larva 
and measures 8 to 9 mm long; in a dorsal 
view, its shape is pyriform; the anterior por- 
tion is highly convex dorsally, becoming 
narrower and cone shaped posteriorly. Ven- 
trally the puparium is flat posteriorly and 
slightly concave anteriorly. The surface is 


776 


coarse, without tubercules. The posterior 
breathing tube is very short and black. 

We reared tristani from July Ist to Oc- 
tober 13, 1993 and from June 20 to October 
10, 1994, at INBio headquarters in Santo 
Domingo, Heredia Province, Costa Rica, at 
an elevation of 1,100 m. During the first 
season we did not successfully rear a single 
adult, just parasitic wasps (Hymenoptera: 
Pteromalidae and Encyrtidae). Larvae and 
puparia were found from June to October, 
during the rainy season in Costa Rica, when 
humidity favors scale insect development. 
Adults were present from February to Au- 
gust. 

We found tristani larvae feeding on scale 
insects (Homoptera: Sternorrhyncha: Coc- 
coidea: Coccidae) on various plant species 
(Rutaceae: Citrus sp., Fabaceae: Acacia sp., 
Gliricidia sepium (Jacq.) Kunth ex Walp, 
Solanaceae: Datura sp., Rubiaceae: Coffea 
arabica Linnaeus, Lamiaceae: Ocimum 
basilicum Linnaeus). 

The larvae use a glue-like saliva (Roth- 
eray 1986) to defend themself against ag- 
gressive organisms such as ants. Two Cam- 
ponotus ants were seen attacking a larva 
that responded by turning the anterior por- 
tion of the body towards the ant and re- 
leasing a drop of this liquid substance on 
them. The ants were trapped for at least 2 
minutes before freeing themselves. 

Several species of parasitic wasps were 
found to attack tristani, all of them emerg- 
ing from puparia. All reared parasitic wasps 
belong to the Encyrtidae, mainly Syrpho- 
phagus sp. which attacks the larval stage 
(Noyes 1995), and Pteromalidae, mainly 
Pachyneuron sp. In most cases, several to 
many (4 to 75 encyrtids, 1 to 33 pterom- 
alids) emerged from a single puparium. 
These natural enemies seem to be very ef- 
fective against tristani larvae. Neither Pa- 
chyneuron nor Syrphophagus are host spe- 
cific in Ocyptamus, both are hyperparasi- 
toids in aphids (Homoptera: Aphididae) or 
primary parasitoids in syrphids (Hanson 
and Gauld 1995). Syrphophagus also para- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


sitizes Braconidae and Aphelinidae hyme- 
nopterans (Noyes 1995). 

In our first attempts to rear the fly (60 
puparia), we got only parasitic wasps (636 
Encyrtidae and 260 Pteromalidae). So we 
started taking very recently formed puparia 
(17) and were then successful in getting 
adult flies (12). This suggests that female 
wasps are ovipositing on freshly formed pu- 
paria, perhaps attracted by the chemicals re- 
leased during the transformation into pupae. 


KEY TO SPECIES OF OCYPTAMUS 
(MIMOCALLA) 


1. Legs black; mesonotum black; wing broadly 
black anteriorly; large, body length 20 mm . . 
A seenta co: Circe ctekn aioe here mene Sb sc giganteus 
— Legs partially or entirely pale, always with me- 
sotibia yellow; wing hyaline or at most narrow- 
ly light brownish anteriorly 
. Metafemur and tibia black; mesonotum black; 
face yellow; large, body length 16 mm or more 
FRR eR ee in arg coro Bnala o-oo & willistoni 
— Metafemur and tibia partially pale, brownish 
orange to yellow; mesonotum at least yellow 
on postalar callus; if mesonotum extensively 
dark, then face extensively dark; smaller, body 
length less than 16mm -:--. °° 2232. 3 
3. Abdomen yellow on Ist, 2nd and basal 4% of 
3rd segments, elsewhere black; pleuron entire- 
ly yellow; legs yellow. Alula microtrichose . . 
Seer 2 ged aa Oe, AY Ree nymphaea 
— Abdomen not as such, with Ist and 2nd seg- 
ment partially dark, brown to black; pleuron 
and legs partially dark 
4. Postpronotum brownish black, only margins 
pale; notopleuron extensively brownish black; 
face extensively brownish black, only narrowly 
yellow laterally; abdomen appearing totally 
dark, abdominal pale fasciae greatly reduced 
SRP ee PROTA Lae chars Co cof erebus 
— Postpronotum and notopleuron extensively yel- 
low; face yellow or with narrow brown vitta; 
abdomen with distinct yellow fasciae 
5. Alula almost entirely bare. Antenna pale, or- 
ange to brownish orange; scutellum yellow or 
obscurely grayish on disc (West Indies) 
a age a gailcy Dea = ita ks Ae COR capitatus 
— Alula more extensively microtrichose, entirely 
(2) or partially bare medially (d). Antenna and 
scutellum usually partially dark (not West In- 
dies) 
6. 2nd tergum brown to black, except broadly yel- 
low basolaterally and with medial triangular 
macula; scutum with lateral yellow vitta con- 
tinuous behind transverse suture; abdominal 


WN 


VOLUME 102, NUMBER 4 


fasciae on 3rd & 4th terga broad, about % as 

Wi GeraSpSeCIMenterps vas seri ase « Alghct ce lees tristani 
— 2nd tergum reddish brown except narrowly 

yellow basolaterally; scutum with lateral yel- 

low vitta broadly interrupted behind transverse 

suture; abdominal fasciae on 3rd & 4th terga 

narrow, about '/, as wide as segment..... 


Ocyptamus (Mimocalla) bonariensis 
(Bréthes) 
(Figs. 2—4) 


Baccha bonariensis Bréthes 1905: 340 Ar- 
gentina, Buenos Aires (HT ¢6 MACN 
lost); Kertész 1910: 157 (cat. cit.); Fluke 
1957-5 153.(Cat:. cit.). 

Ocyptamus bonariensis: Thompson et al. 
19762 13 .(Cat:-cit.). 

Salpingogaster flukei Curran 1941: 284 Ec- 
uador, Tungurahua, Banos (HT 4 
AMNH); Arnaud and Owen 1981: 125 
(type data). New synonym. 

Baccha flukei: Hull 1949: 98, 105 (key ref., 
note); Fluke 1956: 246 (cat. cit.). 

Ocyptamus flukei: Thompson et al. 1976: 
18 (cat. cit.). 

Baccha (Mimocalla) phobia Hull 1943b: 51 
Ecuador, Tungurahua, Banos, Chaupi 
(HT ¢ AMNH); Hull 1949: 240 (fig. (ab- 
domen pattern)). Syn. Hull 1949: 98. 

Baccha (Mimocalla) polista Hull 1943c: 89 
Brazil, Santa Catarina, Nova Teutonia 
(HT 56 AMNH); Hull 1949: 98, 105, 236 
(fig. (abdomen pattern), key ref.). New 
synonym. 

Baccha nigriventris (misidentifications): 
Sack 1920: 249, 1921: 135 (puparium); 
Bruch 1923: 1 (cat. cit.); Knutson 1971: 
29 (notes). 


Male.—Head: Face yellow, with light 
brown medial vitta, sparsely white polli- 
nose laterally, only densely pollinose nar- 
rowly along eye margin, white pilose ex- 
cept black pilose ventrad to antenna; gena 
yellow, shiny, bare; lunule yellow except 
brown medially; frontal triangle brown on 
anterior 42, yellow posteriorly, black pilose; 
vertical triangle black, black pilose; occiput 
black except yellow on ventral ¥, densely 
whitish gold pollinose ventrally becoming 


TY 


more golden dorsally, white pilose v« \trally 
becoming more golden dorsally; anicnna 
brownish black except reddish brown ba- 
soventrally on basoflagellomere, black pi- 
lose; basoflagellomere elongate, about % 
longer than wide. 

Thorax: Mainly brownish black with 
yellow maculae; postpronotum yellow, with 
rest of prothorax brownish yellow; scutum 
black except with broad yellow vitta run- 
ning from postpronotum to and including 
anterior % of postalar callus but broadly in- 
terrupted behind transverse suture and dor- 
sad to wing base, with vitta about as wide 
as postpronotum, with black areas black 
pollinose except for broad golden pollinose 
submedial vitta which tapers posteriorly 
and does not reach scutellum and a very 
narrow indistinct brown pollinose medial 
vitta on anterior % which expands into a 
broader golden pollinose vitta at level of 
postalar callus, long yellow pilose anteri- 
orly and laterally, short intermixed black 
and yellow pile elsewhere; scutellum 
brownish black except yellow base and api- 
cal margin, short sparse black pilose with a 
few intermixed yellow pili basally; subscu- 
tellar fringe singular, white; pleuron black- 
ish brown except yellow dorsad to procoxa 
and broadly yellow on posterior anepister- 
num, dorsal % of katepisternum, anterior %2 
anepimeron and anterior % katatergum, 
very sparsely white pollinose except denser 
dorsally on katepisternum, yellow pilose; 
plumula very short, white; calypter yellow- 
ish white except dorsal margin brownish; 
halter brownish yellow. Legs: Procoxa or- 
ange, sparsely gray pollinose, yellow pi- 
lose; mesocoxa brown, black pilose medi- 
ally, yellow pilose laterally; metacoxa 
black, long black pilose with a few inter- 
mixed yellow pili laterally; protrochanter 
orange; mesotrochanter brownish black; 
metatrochanter black, long black pilose; 
profemur orange, yellow pilose; mesofemur 
orange except blackish basally and apico- 
posteriorly, yellow pilose except for a few 
black pili on dorsoposterior surface; meta- 
femur orange except black ventrally, orange 


778 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


| / yy i 
WHat 


Figs. 2-7. Male genitalia. 2, Ocyptamus bonariensis, 9th sternum, aedeagus and associated structures, lateral 
view. 3. O. bonariensis, 4th sternum, ventral view. 4, O. bonariensis, 9th tergum and associated structures, 
lateral view. 5, O. tristani, 9th sternum and associated structures, lateral view. 6, O. tristani, 4th sternum, ventral 
view. 7, O. tristani, 9th tergum and associated structures, lateral view. 


VOLUME 102, NUMBER 4 


pilose except black pilose ventrally; pro and 
mesotibiae and tarsi yellow, yellow pilose; 
metatibia orange except yellow basal % and 
apex, yellow pilose; metatarsus brownish 
orange, yellow pilose. Wing: Narrowly 
brown anteriorly, hyaline posteriorly, mi- 
crotrichose except bare cell R1 anterior to 
vein Rs, cell R, cell BM, anterobasal % of 
cell CuP, cell R4+5 posterior to spurious 
vein, narrowly on basoposteriorly in cell 
DM, basomedial % of cell CuA1, anterior 
to vein A2; alula entirely microtrichose to 
bare basomedially, normal, as wide as cell 
CuP; brown area including base and ex- 
tending posteriorly to veins R and R1. 
Abdomen: Dark brown with yellow fas- 
ciae; 1st tergum brownish black except yel- 
low basally, white pilose basolaterally, 
black pilose elswhere; 2nd tergum brownish 
black except narrowly yellow basolaterally, 
yellow pilose basolaterally, black pilose 
elsewhere; 3rd tergum yellow on basal \, 
brownish black apically, yellow pilose on 
yellow area, black pilose elsewhere; 4th ter- 
gum brown except yellow basal ;, black 
pilose except yellow pilose basolaterally; 
5th tergum brown, black pilose except yel- 
low laterally; genitalia brown and yellow, 
black pilose; 1st sternum brown, white pi- 
lose; 2nd sternum brown except yellow bas- 
al %, yellow pilose; 3rd & 4th sterna brown 
except narrowly yellow basally, black pi- 
lose; 5th sternum brown, yellow pilose. 
Male genitalia: See Figs. 2—4. 
Female.—Similar to male except for nor- 
mal sexual dimorphism and frons black an- 
teriorly except broadly yellow along eye 
margin, brown on medial %, black posteri- 
orly, shiny on anterior %, sparsely gray pol- 
linose medially, black pollinose on poste- 
rior %4, short black pilose. 
Length.—15 mm; wing, 12 mm. 
Distribution.—Ecuador*, Peru*, Brazil 
(Santa Catarina*), Uruguay*, Argentina 
(Buenos Aires*, Misiones*). 
Types.—Baccha bonariensis was de- 
scribed from a unique male from Palermo, 
Buenos Aires, Argentina, collected on 11 
April 1904. A search of the Diptera collec- 


7719 


tion of MACN failed to find this spc -imen. 
However, there is a single reared female 
from Buenos Aires (9 Dec 1904) which was 
identified by Bréthes. 

Salpingogaster flukei was described from 
a series of 4 males from Ecuador collected 
by W. Clark McIntyre. The holotype is now 
in the AMNH and was examined. Paratypes 
are in the CNC and UWEC. 

Baccha phobia was described from a se- 
ries of 2 males from Ecuador collected by 
W. Clark McIntyre. The holotype is now in 
the AMNH and paratype is in the CNC. 
Both were examined. 

Baccha polista was described from a se- 
ries of 6 specimens from Brazil (Santa Ca- 
tarina) collected by Fritz Plaumann and in 
the Fluke Collection. The holotype is now 
in the AMNH and was examined. Paratypes 
are in the CNC and UWEC. 

Material examined (15 ¢, 6 2).—EC- 
UADOR. Types of flukei and phobia. 
PERU. Cuzco, Kilometer 94, Machu Picchu 
RailRoad, 7 Mar 1978, PB M. Marsh (¢ 
USNM ENT 00037937 USNM); Lima, 
Bosque Carrion, 2,400 m, 16 April 1987, P. 
Hocking (6 USNM ENT 00030147 
USNM). BRAZIL. Santa Catarina: types of 
polista; Nova Teutonia, 300-500 m, Jan 
1965, E Plaumann (26 2 USNM ENT 
00037924-6 USNM); ... Feb 1965 (¢ 
USNM ENT 00037927 USNM); ... Feb 
1972 (6 USNM ENT 00037930 USNM); 
... Nov 1971 (6 USNM ENT 00037929 
USNM); ... Feb 1972, E Plaumann (¢d 
USNM ENT 00037977 USNM); ... Dec 
1964 (6 USNM ENT 00037928 USNM). 
URUGUAY. Montevideo, “Rec. 3.1.42,” 
H. L. Parker (2 with puparium USNM 
ENT 00037938 USNM). ARGENTINA. 
Buenos Aires: Buenos Aires, 4 Dec 1904 
(MACN); Moreno, Dec 1971, Fritz (6 
USNM ENT 00037931 USNM); ... Dec 
1972 (2 2 USNM ENT 00037932-3 
USNM); ... Jan 1973 (2 USNM ENT 
00037934 USNM); Villa Elisa, 15—29 Dec 
1979,°G; My & O: S: FlinteIr (2 USNM 
ENT 00037935 USNM); Santa Catalina, 
near Buenos Aires, 10 Nov 1970, J. W. 


780 


Boyes (6 USNM ENT 00022541 CNC). 
Misiones: Dos de Mayo, Dec 1973, M. 
Fritz (6 USNM ENT 00037936 USNM). 

Discussion.—Ocyptamus bonariensis 1s 
very similar to erebus, but differs in having 
distinct yellow pollinose vittae on the me- 
sonotum and broader yellow fasciae on the 
abdomen. The shape of the male cercus is 
also distinctive. 

The concepts of flukei Curran and polista 
Hull are very close. Hull (1949: 105) sep- 
arated them by two characters (shape of the 
2nd tergum (2nd tergum narrow, only as 
wide basally as apically (flukei) versus 
broader, about twice as wide apically as ba- 
sally (polista)) and color of hind femur 
(hind femur entirely brownish yellow (flu- 
kei) versus black ventrally (polista))). 
While these differences are supported by 
the material we have examined, we consid- 
er them to be a function of geographic var- 
iation which will be bridged once material 
from intermediate localities are studied. 
Hence, we have synonymized the two con- 
cepts. The name bonariensis was unknown 
to Hull, hence he could not have known 
that it was an older synonym for his polista. 
Also, we include here a single male from 
Peru (Cuzco) with a bare alula, no basal 
yellow fascia on 4th tergum, greatly re- 
duced yellow fascia on 3rd tergum and 
which is slightly larger than typical Mimo- 
calla (16 mm body length, 14 mm wing 
length). As the male genitalia are the same 
as those of typical specimens, we consider 
this to be an aberrant specimen. 


Ocyptamus (Mimocalla) capitatus (Loew) 


Baccha capitata Loew 1863: 14 Cuba (LT 
3 MCZ here designated); Williston 1887: 
124 (Cuba, redescr.); Aldrich 1905: 354 
(cat. Git.): Kertész 1910:.157 (cat: \cit.); 
Fluke 1956: 239 (cat. cit.); Wolcott 1923: 
217, 1936: 346, 1948: 461 (Puerto Rico, 
with Saissetia hemisphaerica on coffee). 

Ocyptamus captitatus: Thompson et al. 
1976: 14; Thompson 1981: 44 (figs. (ab- 
domen pattern, male genitalia, wing), 
West Indies). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Doros insularis Bigot 1883: 330 Cuba (HT 
2 BMNH); Williston 1887: 299 (note). 
Syn. Thompson 1981: 44. 

Xanthogramma insulare: Aldrich 1905: 369 
(cat. cit.); Kertész 1910: 152 (cat. cit.). 

Ocyptamus insularis: Thompson et al. 
1976: 20 (cat. cit.). 

Baccha carlota Curran 1929: 491 Cuba, 
Trinidad Mountain, Mina Carlota (HT @ 
AMNH); Hull 1949: 98, 105. 238, 240 
(fig. (abdomen pattern), key ref.); Fluke 
1956: 239 (cat. cit.); Arnaud and Owen 
1981: 83 (type data). Syn. Thompson 
1981: 44. 

Ocyptamus carlota: Thompson et al. 1976: 
14 (cat. cit.). 


Male.—Head: Face yellow, with light 
brown medial vitta, sparsely white polli- 
nose laterally, only densely pollinose nar- 
rowly along eye margin, white pilose ex- 
cept black pilose ventrad to antenna; gena 
yellow, shiny, bare; lunule brown except 
yellow submedially; frontal triangle yellow 
except brown along lunule, black pilose; 
vertical triangle black, black pilose; occiput 
black except yellow on ventral ’;, densely 
golden pollinose becoming more brownish 
on dorsal %, yellow pilose except pile more 
golden dorsally; antenna orange except 
more brownish orange on dorsoapical %4 of 
basoflagellomere, black pilose; basoflagel- 
lomere elongate, slightly less than twice as 
long as wide. 

Thorax: Mainly black with yellow mac- 
ulae; postpronotum yellow; scutum black 
except with broad yellow vitta running 
from postpronotum to and including ante- 
rior % of postalar callus but broadly inter- 
rupted behind transverse suture and dorsad 
to wing base, with vitta about as wide as 
postpronotum, with black areas black pol- 
linose except for broad golden pollinose 
submedial vitta which tapers posteriorly 
and does not reach scutellum and a very 
narrow indistinct brown pollinose medial 
vitta on anterior 7% which expands into a 
broader golden pollinose vitta at level of 
postalar callus, long yellow pilose anteri- 


781 


VOLUME 102, NUMBER 4 


hin ‘i 
Wr 
a iS 
‘ Ww 
5 
feet 
og 


a, Ab 12 


Male genitalia. 8, Ocyptamus giganteus, 9th sternum, aedeagus and associated structures, lateral 


Figs. 8-13. italia. 8, ) é 
view. 9, O. giganteus, 4th sternum, ventral view. 10, O. giganteus, 9th tergum and associated structures, lateral 
view. 11, O. nymphaea, 9th sternum and associated structures, lateral view. 12. O. nymphaea, 4th sternum 


ventral view. 13, O. nymphaea, 9th tergum and associated structures, lateral view 


782 


orly and laterally, short intermixed black 
and yellow pile elsewhere; scutellum yel- 
low except disc blackish, short sparse black 
pilose with a few intermixed yellow pili ba- 
sally; subscutellar fringe singular, white; 
pleuron black except yellow dorsad to pro- 
coxa and broadly yellow on posterior ane- 
pisternum, dorsal % of katepisternum and 
anterior % anepimeron, very sparsely white 
pollinose except denser dorsally on katepis- 
ternum, yellow pilose; plumula very short, 
yellow; calypter yellowish white except 
dorsal margin brownish; halter yellow. 
Legs: Procoxa brownish black except yel- 
low apex, sparsely gray pollinose, yellow 
pilose; mesocoxa black, black pilose me- 
dially, yellow pilose laterally; metacoxa 
black, long black pilose with a few inter- 
mixed yellow pili laterally; protrochanter 
yellowish brown; mesotrochanter brownish 
black; metatrochanter black, long black pi- 
lose; profemur yellow, yellow pilose; me- 
sofemur yellow except black on basodorsal 
¥%, yellow pilose except for a row of black 
pili on dorsoposterior surface; metafemur 
black except yellow apical ¥,, black pilose 
except yellow pilose on yellow area; pro 
and mesotibiae and tarsi yellow, yellow pi- 
lose; metatibia black except yellow basal % 
and apex, black pilose; metatarsus yellow, 
yellow pilose. Wing: Brown anteriorly, hy- 
aline posteriorly, microtrichose except bare 
cell R posterior to spurious vein on apical 
¥%, cell BM, anterobasal % of cell CuP, cell 
R4+5 posterior to spurious vein, narrowly 
on basoposteriorly in cell DM, basomedial 
¥% of cell CuA1, anterior to vein A2; alula 
bare, normal, as wide as cell CuP; brown 
areas including base and extending poste- 
riorly to vein M to level of end of vein RS, 
to spurious vein in apical portion of cell R, 
to vein R4+5 except narrowly hyaline 
along sinuate portion. 

Abdomen: Dark brown with yellow fas- 
ciae and orange apex; Ist tergum yellow ex- 
cept brownish black apicomedial %, yellow 
pilose; 2nd tergum brownish black except 
for yellow basolateral triangular macula, 
with maculae broadly separated medially, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


yellow pilose basolaterally, black pilose 
elsewhere; 3rd tergum yellow on basal %, 
brownish black apically, yellow pilose on 
yellow area, black pilose elsewhere; 4th ter- 
gum brown except with yellow basal fascia 
which is narrowly isolated from lateral mar- 
gin and occupies basal %4, with apical por- 
tion of tergum becoming more orange, 
black pilose; 5th tergum orange, black pi- 
lose; genitalia yellowish orange, black pi- 
lose; 1st sternum yellow, yellow pilose; 2nd 
sternum brown except yellow basally, yel- 
low pilose; 3rd sternum brown except yel- 
low basal %, black pilose except yellow pi- 
lose basally; 4th sternum brown, brown pi- 
lose; 5th sternum orange except yellow api- 
cally, brown pilose. Male genitalia. See 
Figs. 56a—c in Thompson (1981: 59); es- 
sentially the same as tristani (Figs. 5—7). 

Female.—Similar to male except for nor- 
mal sexual dimorphism and frons yellow 
except may be darker medially and along 
lunule, orange pollinose except maybe 
more brownish medially, short black pilose. 

Length.—14 mm; wing, 11 mm. 

Variation.—The brownish black color in 
some examined individuals is brownish or- 
ange; also the antenna may be entirely or- 
ange and scutellum entirely yellow. 

Distribution.—West Indies (Cuba*, Do- 
minican Republic*, Puerto Rico*, Lesser 
Antilles*). 

Types.—Baccha capitata was described 
from an unspecified number of male spec- 
imens from Cuba from Gundlach. In the 
MCZ, there is a male with a silver square 
and labelled ‘‘capitata mihi’? in Loew’s 
hand. This specimen is here designated lec- 
totype so as to fix and stabilize the current 
concept of the name, and has been so la- 
belled. 

Doros insularis was described fron a 
unique female specimen from Cuba. The 
holotype is now in the BMNH and has been 
examined. 

Baccha carlota was described from a 
unique female from Cuba collected by G. 
Salt and in the Curran collection. The ho- 


VOLUME 102, NUMBER 4 


lotype is now in the AMNH and was ex- 
amined. 

Material examined (8 3, 7 2).—CUBA. 
Types of capitata, insularis & carlota. DO- 
MINICAN REPUBLIC. Pedernales, 23.5 
km N Cabo Rojo, 18-16N 71-38W, 540 m, 
13 July 1990, C. Young, J. Rawlins & S. 
Thompson (6 USNM ENT 00037907 
USNM). PUERTO RICO. Comerio, 10 
Aug 1913, J. R. Johnston (6 USNM ENT 
00037900 USNM); Mayaguez, 4 Mar 1912, 
C. W. Hooker (2 USNM ENT 00037901 
USNM); ... 26 Jul 1915, R. H. van Zwa- 
lenberg (6 USNM ENT 00037902 
WSNM); .... > May’ 1939;H. K.\Plank.G 
66 2 22 with puparia USNM ENT 
00037904-6, ... 72-73 USNM) VIRGIN 
ISLANDS. St. Croix, 6 Jul 1963, R. M. 
Bond & R. Boyd (¢6 % USNM ENT 
00037899, ... 903 USNM). 

Discussion.—Ocyptamus capitatus is 
similar to tristani and bonariensis in having 
distinctive yellow pollinose vittae on me- 
sonotum, but differs from these species in 
having a yellow scutellum and the apex of 
the abdomen orange to brownish orange. 


Ocyptamus (Mimocalla) erebus (Hull) 
(Figs. 14—16) 


Baccha (Mimocalla) erebus Hull 1943d: 
54; 1943e: 137 Brazil, Santa Catarina, 
Nova Teutonia (HT ¢ AMNH); Hull 
1949: 98, 105, 240 (fig. (abdomen pat- 
tern), key); Fluke 1956: 245 (cat. cit.). 

Ocyptamus erebus: Thompson et al. 1976: 
17 (cat: Cit.). 


Male.—Head: Face yellowish brown, 
with dark brown medial vitta, sparsely 
white pollinose laterally, only densely pol- 
linose narrowly along eye margin, white pi- 
lose except black pilose ventrad to antenna; 
gena yellow, shiny, bare; lunule black ex- 
cept brownish submedially; frontal triangle 
black except narrowly yellow along eye 
margin, shiny except dull pollinose on pos- 
terior %, black pilose; vertical triangle 
black, black pilose; occiput black except 
yellow on ventral Y,, densely gray pollinose, 


783 


white pilose; antenna black excepi more 
brownish orange basoventrally on basofia- 
gellomere, black pilose; basoflagellomere 
elongate, slightly less than twice as long as 
wide. 

Thorax: Mainly brownish black; post- 
pronotum reddish brown; scutum black ex- 
cept with obscure narrow reddish brown 
vitta posterior to postpronotum and another 
anterior to and including anterior % of pos- 
talar callus, with area behind transverse su- 
ture and dorsad to wing base dark, with 
black areas black pollinose except for nar- 
row golden pollinose submedial vitta which 
tapers posteriorly and does not reach scu- 
tellum and a narrow golden pollinose me- 
dial vitta on posterior % at level of postalar 
callus, long yellow pilose anteriorly and lat- 
erally, short intermixed black and yellow 
pile elsewhere; scutellum dark brown ex- 
cept brownish yellow on base and apical 
margin, short sparse black and yellow pi- 
lose; subscutellar fringe singular, white; 
pleuron black except obscurely yellow on 
posterior anepisternum, dorsal % of katepis- 
ternum and medially on katatergum, very 
sparsely white pollinose except denser dor- 
sally on katepisternum, white pilose; plu- 
mula very short, yellow; calypter brownish 
white except dorsal margin brown; halter 
brown. Legs: Procoxa yellow, sparsely 
white pollinose, yellow pilose; mesocoxa 
orange, brownish-orange pilose; metacoxa 
black, long black pilose with a few inter- 
mixed yellow pili laterally; pro and meso- 
trochanter brown; metatrochanter black, 
long black pilose; pro and mesofemur red- 
dish brown, yellow pilose except for a few 
black pili on dorsoposterior surface; meta- 
femur reddish brown except yellow apical 
/;, Orange pilose except black apicoventral- 
ly; pro and mesotibiae and tarsi yellow, yel- 
low pilose; metatibia yellow except reddish 
brown medially, orange pilose; metatarsus 
dark brown, yellow pilose. Wing: Light 
brown anteriorly, hyaline posteriorly, mi- 
crotrichose except bare cell R1 anterior to 
vein Rs, cell R, cell BM, anterobasal % of 
cell CuP, cell R4+5 posterior to spurious 


784 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 14-19. Male genitalia. 14, Ocyptamus erebus, 9th sternum, aedeagus and associated structures, lateral 
view. 15, O. erebus, 4th sternum, ventral view. 16, O. erebus, 9th tergum and associated structures, lateral view. 
17, O. willistoni, 9th sternum and associated structures, lateral view. 18, O. willistoni, 4th sternum, ventral view. 
19, O. willistoni, 9th tergum and associated structures, lateral view. 


VOLUME 102, NUMBER 4 


vein, narrowly on basoposteriorly in cell 
DM, basomedial % of cell CuA1, anterior 
to vein A2; alula microtrichose, normal, as 
wide as cell CuP; brown areas including 
base and extending posteriorly to vein R 
and R1. 

Abdomen: Dark brown with greatly re- 
duced yellow fasciae; Ist tergum brownish 
black, white pilose; 2nd tergum brownish 
black except for small yellow basolateral 
triangular macula, with maculae broadly 
separated medially, yellow pilose basolater- 
ally, black pilose elsewhere; 3rd and 4th 
terga brownish black except for very nar- 
row obscure yellow fasciate maculae broad- 
ly separated medially and occupying less /,, 
of tergal length, yellow pilose laterally, 
black pilose elsewhere; 5th tergum black, 
black pilose; genitalia brownish black, 
black pilose; Ist sternum brown, white pi- 
lose; 2nd sternum brown except yellow ba- 
sally, yellow pilose; 3rd and 4th sterna 
brown except yellow apical margin, black 
pilose; 4th sternum brown, brown pilose; 
5th sternum brownish orange, brown pilose. 
Male genitalia. See Figs. 14-16. 

Female.—Similar to male except for nor- 
mal sexual dimorphism and frons black ex- 
cept narrow yellow vitta along eye margin, 
shiny anteriorly, gray pollinose on medial 
¥%, black pollinose posteriorly, short black 
pilose. 

Length.—13 mm; wing, 11 mm. 

Distribution.—Brazil (Minas Gerais*, 
Sao Paulo*, Santa Catarina*). 

Type.—Baccha erebus was described 
from 4 male specimens collected in south- 
ern Brazil (Santa Catarina) by Fritz Plau- 
mann and in the Fluke Collection. The ho- 
lotype is now in the AMNH and was ex- 
amined. Paratypes are in the CNC and the 
University of Wisconsin Entomology Col- 
lection. The name was first validated in a 
key which appeared before the original de- 
scription. 

Material examined (10 ¢d 6 ?).—BRA- 
ZIL. Minas Gerais: Arceburgo, FE Fortaleza, 
Dec 1946, Barretto (2 22 USNM ENT 
00037974—5 MZUSP). Sao Paulo: Cantar- 


785 


eira, Chapadao, Nov 1946, Barreiic (2 
USNM ENT 00037976 MZUSP). Santa “a- 
tarina: types of erebus; Nova Teutonia, 27- 
11S 52-23W, 300-500 m, Feb 1965, FE 
Plaumann (2 66 USNM ENT 00037908— 
9 USNM); ... Feb 1972 (2 USNM ENT 
00037916 USNM); > Mar 21957, (2 
USNM ENT 00037910 USNM); ... Mar 
1960 (2 USNM ENT 00037912 USNM); 
... Nov 1952 (6 USNM ENT 00037911 
USNM); ... Nov 1971 (3 66 USNM ENT 
00037913—5 USNM). 

Discussion.—Ocyptamus erebus is a dark 
version of bonariensis, the yellow vittae on 
the mesonotum and abdominal fasciae are 
greatly reduced, obscure or absent. The 
shape of the male cercus is also very dis- 
tinctive. 


Ocyptamus (Mimocalla) giganteus 
(Schiner) 
(Figs. 8-10) 


Baccha gigantea Schiner 1868: 340 *‘South 
America’?> = Venezuela (LT 6 NMW 
here designated); Williston 1886: 314 
(cat. cit.); Kertész 1910: 160 (cat. cit.); 
Fluke 1956: 154 (cat. cit.). 

Ocyptamus giganteus: Thompson et al. 
1976: 19 (cat. cit.). 


Male.—Head: Mainly black; face black, 
with narrow yellow medial vitta, white pol- 
linose except narrowly shiny medially, with 
pollinosity denser along eye margin becom- 
ing sparser medially, black pilose; gena 
black, shiny, bare; frontal triangle black ex- 
cept narrowly yellow medially along eye 
margin, black pilose, black pollinose except 
white pollinose on yellow areas; lunule 
black except brownish submedially; vertical 
triangle black, black pollinose and pilose; 
occiput black, black pilose except mesial 
row of cilia mainly white and white pilose 
on ventral /;; antenna black except slightly 
reddish basoventrally on basoflagellomere, 
black pilose, elongate, with basoflagellom- 
ere about twice as long as wide. 

Thorax: Black, black pilose except white 
pilose along anterior margin of scutum and 


786 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


/ wes i Prt « 
hah 4 ; “a f ( ae 


J » 


LDONZO 


21 


ie 


[ 


A 


¥ 
x 


Saal st eee 
whit st ae ; : ed Dia 


C 
Figs. 20-23. 


LDONZO 


Immature stages of Ocyptamus tristani. 20, Larva, a, lateral view, b, dorsal view, c, ventral 


view. 21, Larva, anterior view. 22, Larval posterior spiracle, posterior view. 23, Puparium, dorsoblique view. 


posterior anepisternum; scutum brownish 
gray to black pollinose, with whitish to gray 
submedial and medial vittae; scutellum 
brownish black pollinose, subscutellar 
fringe singular, white; pleuron black polli- 
nose except katepisternum white pollinose 
on dorsal %; calypter black; plumula long; 


halter black. Legs: Black except grayish 
white basal %4 of mesotibia, black pilose; 
metatrochanter with distinct pile tuft; me- 
tafemur without long pile. Wing: Dark an- 
teriorly, hyaline posteriorly, microtrichose 
except bare on anterobasal % of cell CuP; 
alula dark, normal, as wide as cell CuP, mi- 


VOLUME 102, NUMBER 4 


crotrichose; black area covering base of 
wing including base of cell CuP, all of cell 
BM, most of cell R except posterior to spu- 
rious vein on apical %, anterobasal corner 
of cell R4+5, extending almost to level of 
middle of cell R4+5; brownish orange area 
between black area and gray apex; gray 
area covering apex of wing, anterior to vein 
R4+5, distal to end of vein R1. 

Abdomen: Black except yellow basola- 
teral triangular macula on 2nd tergum and 
narrow yellow fascia on base of 3rd, black 
pilose except white pilose on Ist, bases of 
2nd and 3rd terga and on yellow areas and 
male genitalia. Male genitalia. See Figs. 8— 
10. 

Female.—Similar to male except for nor- 
mal sexual dimorphism and frons black ex- 
cept very narrowly yellow on ventrolateral 
73, black pollinose sublaterally becoming 
gray pollinose medially, black pilose. 

Length.—20 mm; wing, 14 mm. 

Distribution.—Costa Rica*, Colombia™*, 
Venezuela*. 

Types.—Baccha giganteus was described 
from an unspecified number of specimens 
from an unspecified locality in South Amer- 
ica. In NMW, there are a male and two fe- 
males from Venezuela, Lindig collector, la- 
belled as giganteus. The male is here des- 
ignated lectotype so as to fix and stabilize 
the current concept of the name, and has 
been so labelled. 

Material examined (3 6 4 ?).—VENE- 
ZUELA. Types of giganteus. COSTA 
RICA. Puntarenas: Las Alturas, Cerro Chai, 
2,100 m, 14 Aug 1995, T. Pape (2 6 USNM 
ENT 00037918 NRS, 00030146 
USNM); Quijada del Diablo, 3.1 km NE de 
Mellizas, 1,800 m, LS 316900_600600, 17 
Aug 1997, A. Picado, Lot# 47696 (2 IN- 
BIOCRIO02546773 INBIO). COLOMBIA. 
Antioquia: Caldas, Oct 1973, R. Velez (2 
USNM ENT 00037917 USNM). 

Discussion.—Ocyptamus giganteus is 
readily recognized by its overall dark col- 
oration, black thorax, legs, anterior wing 
margin and abdomen except for a single 
yellow fascia on the base of the 3rd tergum. 


787 


Also, true to its name, giganteus is the \arg- 
est species of the genus. 


Ocyptamus (Mimocalla) nymphaea (Hull) 
(Figs. 11-13) 


Baccha nymphaea Hull 1943a: 40 Brazil, 
Sao Paulo, Campinas (HT 6 USNM); 
Hull 1949: 98, 105, 238 (fig. (abdomen 
pattern), key ref.); Fluke 1956: 256 (cat. 
Cit): 

Ocyptamus nymphaea: Thompson et al. 
1976: 23.(Cat. Cit.): 


Male.—Head: Face orange, shiny except 
narrowly white pollinose laterally, white pi- 
lose; gena yellow, shiny, bare; frontal tri- 
angle orange on posterior % along eye mar- 
gins, brownish black anteromedially, black 
pilose, shiny anteriorly, dull pollinose pos- 
teriorly; lunule orange except black medi- 
ally; vertical triangle black, black pilose; 
occiput black on dorsal %, orange basally, 
densely yellow pollinose, yellow pilose; an- 
tenna orange except more brownish orange 
on dorsoapical % of basoflagellomere, 
densely black pilose; basoflagellomere 
elongate, slightly less than twice as long as 
wide. 

Thorax: Orange except brownish black 
medially on scutum; scutum broadly orange 
laterally and anterior to scutellum, brown- 
ish black and brown-black pollinose medi- 
ally, with broad golden pollinose submedial 
vitta and a very narrow indistinct brown 
pollinose medial vitta on anterior % which 
expands into a large triangular golden pol- 
linose macula on posterior 4%, long yellow 
pilose anteriorly and laterally, short inter- 
mixed black and yellow pile elsewhere; 
scutellum orange, short sparse black pilose; 
subscutellar fringe singular, yellow; pleuron 
sparsely white pollinose, yellow pilose; ca- 
lypter orange except dorsal margin brown- 
ish; plumula absent; halter orange. Legs: 
Orange except metatarsus brownish orange, 
yellow pilose except black ventral pile tufts 
on metacoxa and trochanter and scattered 
intermixed black pile on posterior surface 
of mesofemur and medially on metatibia. 


788 


Wing: Hyaline except cell C brownish or- 
ange and cell Sc brownish, extensively mi- 
crotrichose except bare: cell R posterior to 
spurious vein, cell BM, anterobasal % of 
cell CuP, cell R1 anterior to spurious vein; 
cell R4+5 posterior to spurious vein, nar- 
rowly on base of cells DM and CuA1. 

Abdomen: Orange and orange pilose on 
Ist, 2nd and basal %4 of 3rd segment; black 
and black pilose elsewhere. Male genitalia. 
See Figs. 11-13. 

Female.—Similar to male except for nor- 
mal sexual dimorphism and frons black and 
shiny on anterior 4%, orange and orange pol- 
linose on posterior %4, short black pilose. 

Length.—14 mm; wing, 11 mm. 

Distribution.—Brazil (Sao Paulo*, Minas 
Gerais*). 

Type.—Baccha nymphaea Hull was de- 
scribed from two reared males from Brazil. 
Although Hull headed his description as 
‘female,’ the types are males. The holo- 
type is preserved in the USNM and has 
been examined. The paratype is now in 
CNC. 

Material examined (5 6 2 ?).—BRA- 
ZIL. Minas Gerais: Arceburgo, E Fortaleza, 
Dec 1946, Barretto (3 ¢6 2 2 USNM ENT 
00037978—82 USNM MZUSP). Sao Paulo: 
Campinas, 14 Oct 1939, H. E G. Sauer 
(Holotype 6 with puparium USNM ENT 
00037983 USNM); Campininha, Mogi 
Guacu, 23 Oct 1970, J. W. Boyes (¢ 
USNM ENT 00022540 CNC). 

Discussion.—Ocyptamus nymphaea is 
readily recognized by the orange and black 
abdomen. The other species of Mimocalla 
have the abdomen fasciate or mostly dark. 


Ocyptamus (Mimocalla) tristani 
Zumbado, new species 
(Figs. 1, 5—7) 


Male.—Head: Face yellow, with light 
orange medial vitta, sparsely white polli- 
nose laterally, only densely pollinose nar- 
rowly along eye margin, white pilose ex- 
cept black pilose ventrad to antenna; gena 
yellow, shiny, bare; lunule yellow except 
brown medially; frontal triangle yellow ex- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


cept brown along lunule, black pilose; ver- 
tical triangle black, black pilose; occiput 
black except yellow on ventral /;, densely 
yellowish-white pollinose ventrally becom- 
ing more golden dorsally, white pilose ven- 
trally becoming more golden dorsally; an- 
tenna brown except more brownish orange 
basoventrally on basoflagellomere and yel- 
low on inner side of scape and pedicel, 
black pilose; basoflagellomere elongate, 
slightly less than twice as long as wide. 
Thorax: Mainly yellow with black mac- 
ulae; prothorax yellow; scutum black ex- 
cept with broad yellow vitta running from 
postpronotum to and including anterior % 
of postalar callus but narrowly attenuated 
dorsad to wing base, with vitta about as 
wide as postpronotum, with black areas 
black pollinose except for broad golden 
pollinose submedial vitta which tapers pos- 
teriorly and does not reach scutellum and a 
very narrow indistinct brown pollinose me- 
dial vitta on anterior % which expands into 
a broader golden pollinose vitta at level of 
postalar callus, long yellow pilose anteri- 
orly and laterally, short intermixed black 
and yellow pile elsewhere; scutellum yel- 
low except disc blackish, short sparse black 
pilose with a few intermixed yellow pili ba- 
sally; subscutellar fringe singular, white; 
pleuron yellow except black narrowly on 
anterior convex surface of posterior anepis- 
ternum, ventral % of katepisternum, poste- 
riorly on katatergum and anteriorly on an- 
atergum, very sparsely white pollinose ex- 
cept denser dorsally on katepisternum, yel- 
low pilose; plumula absent short, yellow; 
calypter yellowish white except dorsal mar- 
gin brownish; halter yellow, with capitulum 
brownish. Legs: Proleg yellow except 
slightly brownish subapically on posterior 
of femur, sparsely gray pollinose, yellow pi- 
lose; mesoleg yellow except sligthly brown- 
ish on base of and subapically on posterior 
of femur, yellow pilose except for a row of 
black pili on dorsoposterior surface; meta- 
coxa brown, long black pilose with inter- 
mixed yellow pili laterally; metatrochanter 
brown, long black pilose; metafemur yel- 


VOLUME 102, NUMBER 4 


low except black ventrally and posteriorly 
except apically, black pilose except yellow 
pilose on yellow area; metatibia yellow ex- 
cept brownish medial %, yellow pilose; 
metatarsus yellow except brownish-orange 
basotarsomere, yellow pilose. Wing: Hya- 
line except stigma brownish and cell Sc yel- 
lowish orange, microtrichose except bare 
cell R posterior to spurious vein on apical 
¥, cell BM, anterobasal % of cell CuP, cell 
R4+5 posterior to spurious vein, narrowly 
on basoposteriorly in cell DM, basomedial 
¥% of cell CuAl, anterior to vein A2; alula 
microtrichose, normal, as wide as cell CuP. 
Abdomen: Dark brown with broad yel- 
low fasciae; Ist tergum yellow on basal %, 
black apically, yellow pilose basally, black 
pilose apically; 2nd tergum brownish black 
except for large yellow basolateral trian- 
gular macula and medial inverted V-shaped 
macula, with basolateral maculae narrowly 
separated medially and occupying basal % 
of tergal length, with V-shaped macula 
slightly beyond middle of tergum and com- 
pletely surrounded by black areas, yellow 
pilose basolaterally, black pilose elsewhere; 
3rd tergum yellow on basal *%, brownish 
black apically, yellow pilose on yellow 
area, black pilose elsewhere; 4th tergum 
yellow on basal 4%, margin and apicolateral 
corner, elsewhere black, yellow pilose; 5th 
tergum yellow except with broad medial 
black fasciate macula, black pilose; geni- 
talia yellow, black pilose; Ist sternum yel- 
low, yellow pilose; 2nd sternum brown ex- 
cept yellow basally, yellow pilose; 3rd ster- 
num brown except yellow basal %, black 
pilose except yellow pilose basally; 4th 
sternum brown, brown pilose; 5th sternum 
orange except yellow apically, brown pi- 
lose. Male genitalia. See Figs. 5—7. 
Female.—Similar to male except for nor- 
mal sexual dimorphism and frons black ex- 
cept yellow broadly along eye margin, short 
black pilose. 
Length.—14 mm, wing, 12 mm. 
Distribution.—Mexico*, El Salvador*, 
Costa Rica*, Colombia*, Venezuela*. 
Holotype.—Male, Costa Rica, Heredia, 


789 


Santo Domingo, Instituto Nacion:! de 
Biodiversidad, 1,100 m, LN 217300 
526200, 2 Jul 1994, M. A. Zumbado, 
voucher# 94-MAZ-28, with puparium 
(INBIOCRI001146854), deposited in IN- 
Bio, Santo Domingo. 

Paratypes (25 6, 25 ?).—MEXICO. 
Vera Cruz: Fortin de la Flores, 9 June 1959, 
H. E. Evans (6 2 USNM ENT00022536-7 
CNC). EL SALVADOR. Santo Tecla, 7 Sep 
L970; lipes’ (Aq i2t) USNMRENE 
00037962—3 USNM). COSTA RICA. Ala- 
juela: Sarchi, 27 Jul 1954, L. A. Salas, ex 
larva feeding on Saisseta hemisphaerica on 
coffee (2 6 2 USNM ENT 00037957-9 
USNM); Upala, Dos Rios, Sect. San Ra- 
mon, 620 m, LN 318100 381900, 18 Mar- 
13 Apr 1995, EF A. Quesada, lot# 5274 (d 
with puparium INBIOCRI002246265 IN- 
BIO). Guanacaste: Volcan Cacao, Cerro 
Pedregal, 1,000 m, Feb—Apr 1989, I. Gauld 
(2 USNM ENT 00037970 MIUCR). He- 
redia: same data as holotype (2 INBI- 
OCRIO01146855 INBIO, 3 with puparium 
INBIOCRIO02153879 INBIO); same local- 
ity as holotype, 4 Jul 1994, M. A. Zumbado 
(36 with puparium, INBIOCRIO02153880 
INBIO); ... 7 July 1994, M. A. Zumbado 
(6 with puparium INBIOCRI002153881 
INBIO); ... Oct 1994, M. A. Zumbado (34 
22 INBIOCRO002153891—5 INBIO); 
11-13 Aug 1994, L. Donzo lot# 3317, (¢ 
INBIOCRIO02112689 INBIO); ... 2 Jul 
1994, J. A. Ugalde (d with puparium, IN- 
BIOCRI002153889 INBIO); ... 4 Jul 
1994, J. A. Ugalde (6 with puparium, IN- 
BIOCRIO02153887 INBIO); 13 Jul 
1994, J. A. Ugalde (2 with puparium, IN- 
BIOCRI002153888 INBIO); Heredia, 1 
Apr 1960, A. Morales, “‘feeding on black 
scale” (6 USNM ENT 00037960 USNM); 
Santo Domingo, 7 Jun 1974, E. Solera (@ 
USNM ENT 00037961 USNM); Belén, San 
Antonio, 950 m, LN 218800 516175, 8 Aug 
1996, M. A. Zumbado, reared from scales 
on Ocimum basilicum (Lamiaceae) (6 2 
with puparia INBIOCRI00215877-8 IN- 
BIO). San José: San José, May 1930, J. E 
Tristan (2 USNM ENT 0037956 USNM); 


790 


San Sebastian, 16 May 1988, A. Retana, 
‘“Comiendo escama”’ (¢ with puparium, 3 
2 USNM ENT 00037964—7 USNM); Cui- 
dad Colén, 800 m, Feb 1990, L. Fournier 
(2 USNM ENT 00037968 USNM); San 
Antonio de Escazt, 1,300 m, Apr 1989, P. 
Hanson (6 USNM ENT 00037969 
MIUCR). COLOMBIA. Antioquia: Medel- 
lin Valley, Apr—Jun 1947, FE L. Gallego (32 
USNM ENT 00037939—41 USNM); . 
[no date], EF L. Gallego, “larva predator of 
aphids’? (2 USNM ENT 00037942 
USNM); Bello, 25 Dec 1970, G. Sanchezg, 
“larva feeding on Saisetia coffeae on or- 
namental palm’ (7 3 5 @ with puparia 
USNM ENT 00037943-54 USNM). VEN- 
EZUELA. Distrito Federal: Cumbre de Bo- 
queron, Frente a Bajo Seco, 1,700 m, 7 Mar 
1970, larva predator of Coccus hesperidum 
on “‘naranjo” (6 with puparium USNM 
ENT 00037955 USNM). 

Etymology.—This species is named af- 
ter José Fidel Tristan Fernandez (1874-— 
1932), the outstanding Costa Rican natu- 
ralist who published the first list of the in- 
sects of Costa Rica (Tristan 1897) and col- 
lected the first specimen of this species 
known to us. 

Discussion.—This species has been 
reared a number of times, and specimens 
have been submitted for identification to 
the Systematic Entomology Laboratory, 
USDA. The species has been reared as a 
predator of various scales (Saissetia cof- 
faea (Walker), S. oleae (Olivier) and Coc- 
cus hesperidum Linnaeus) on a range of 
hosts. The determination labels associated 
with the specimens in the USNM indicate 
that the species was first incorrectly iden- 
tified as Baccha bromleyi by Hull and this 
mistake was perpetuated by subsequent 
specialists (e.g., W. W. Wirth, L. V. Knut- 
son) as either bromleyi or new species near 
bromleyi. Apparently none of these mis- 
identifications were published. 

Ocyptamus tristani is similar to capita- 
tus but differs by the presence of a yellow 
fascia on the 2nd tergum and broader yel- 
low fasciae on the 3rd and 4th terga. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Ocyptamus (Mimocalla) willistoni 
Thompson 
(Figs. 17-19) 


Baccha dolosa Williston 1891: 37 (preocc. 
Walker 1857) AH* H* Mexico, Guer- 
rero, Omiltemi, 8,000 ft.; Amula, 6,000 
ft.; Xucumanatlan, 7,000 ft.; Guatemala, 
Cerro Zunil, 4,000—5,000 ft. (ST 3 
AMNH, BMNH); Aldrich 1905: 355 
(cit.); Kertész 1910: 158 (cit)? al 
1949: 98, 105, 234 (abdomen pattern, 
key ref.); Fluke 1956: 244 (cit.). 

Ocyptamus willistoni Thompson in 
Thompson et al. 1976: 30 (new name for 
dolosa Williston). 


Male.—Head: Face yellow, white pol- 
linose laterally, shiny medially, white pi- 
lose; gena yellow, shiny, bare; frontal tri- 
angle broadly yellow along eye margin, 
black medially, black pilose, sparsely gray 
pollinose medially; lunule yellow except 
black medially; vertical triangle black, 
black pilose; occiput black, densely white 
pollinose, white pilose; antenna black ex- 
cept reddish basally on scape and basofla- 
gellomere; basoflagellomere about % lon- 
ger than wide. 

Thorax: Bluish gray to black except 
yellow base of scutellum and pre-alar area; 
scutum grayish-white pollinose except 
with black pollinose submedial and sub- 
lateral vittae, long white pilose along an- 
terior margin and laterally, short black pi- 
lose elsewhere; scutellum yellow on basal 
%, black elsewhere, short black pilose; 
subscutellar fringe multiple, white; pleuron 
sparsely silver pollinose except dense sil- 
ver pollinose on dorsal % of katepisternum 
and posterior anepisternum, white pilose 
except black pilose on anepimeron; calyp- 
ter white except black dorsal margin; plu- 
mula short; halter yellow. Legs: Coxae 
and trochanters black, silver pollinose, 
black pilose except white pilose apicola- 
terally; pro and mesofemora black except 
yellow apex, white pilose except black pi- 
lose on anteroapical %; metafemur black, 
white pilose except black pilose on apical 


VOLUME 102, NUMBER 4 


¥%; pro and mesotibiae yellowish white, 
white pilose; metatibia black, black pilose; 
pro and meso tarsi black except brownish 
basal % of mesobasitarsomere, white pilose 
on basotarsomere and basally on 2nd tar- 
somere, black pilose elsewhere; metatarsus 
yellowish white to orange on basal 3 tar- 
someres, black apically, white pilose; pile 
tufts distinct on metacoxa and trochanter. 
Wing: Hyaline except black apex and cell 
Sc, extensively bare; microtrichose on api- 
calstaror cell Cy celluSc)apical’/.of cell 
R1, apicomedial % of cell R2+3, apico- 
medial %4 of cell R4+5 and DM, apico- 
medial % of cell CuAl, apicoposterior “4 
of cell CuP, along posterior edge from api- 
cal % of cell CuP to apex; dark apex ex- 
tending apically from end of vein R1 and 
posterior to vein R4+5; alula hyaline, 
bare, normal, as wide as cell CuP. 

Abdomen: \st segement black, white pi- 
lose, sparsely grayish pollinose; 2nd seg- 
ment black except yellow basolaterally, 
black pilose except white pilose basolater- 
ally; 3rd segment yellow on basolateral %, 
black apically, shiny, black pilose on dark 
areas, white pilose on yellow areas; 4th 
segment black, shiny, black pilose; 5th 
segment red except black basomedially on 
tergum, black pilose; genitalia red, yellow 
pilose, see Figs. 17-19. 

Female.—Similar to male except for 
normal sexual dimorphism and: frons yel- 
low on lateral %4, black elsewhere, sparsely 
gray pollinose medially, black pilose; wing 
with black area more extensive, including 
all of cell C and RI. 

Length.—16—17 mm; wing, 12—14 mm. 

Types.—Baccha dolosa was described 
from 5 specimens collected in southern 
Mexico (Guerrero) by H. H. Smith, and 
Guatemala by G. C. Champion. A male 
specimen labelled ‘‘Omilteme, Guerrero, 
8,000 ft., July, H. H. Smith,”’ ‘“‘Biol. Centr. 
Amer., Dipt.—Syrphidae, E D. Godman, 
O. Salvin, 1903-—51,’’ [yellow cotype la- 
bel] and printed Williston determination 
label, in the BMNH is here designated lec- 
totype to fix and stabilize the current con- 


791 


cept of the name, and has been so labelled. 
A paralectotype is in the AMNH. 

Distribution.—Mexico (Guerrero*, Mo- 
relos*, Mexico*, Chiapas*), Guatemala 
(Williston), Costa Rica*. 

Material examined (19 6,8 2 ).—MEX- 
ICO. [no data] (2 with puparium USNM 
ENT 00037920 USNM); “‘Cuernavaca, 8/ 
24” [handwritten, illegible], G. Lassman 
(2 USNM ENT 00022539 CNC). Chiapas: 
Teopisca, 2 miles W of, 4 Aug 1957, J. A. 
Chemsak & B. J. Rannells (6¢6 32 USNM 
ENT 00022547—-55 CNC). Distrito Feder- 
al: Mexico City, Nov 1916, R. Muller (d 
USNM ENT 00037919 USNM); Carretera 
Pedrevall yAjusco, “Kin 7,56"seo. 1979e 
Butze (6 USNM ENT 00028744 UNAM); 
we ZorOct 1979. J Butze, Ie Gutierrez (2 
36 USNM ENT 00028745, 7 UNAM). 
Guerrero: Tuxpan, Torre microondas, 
1,650 m, 16 Aug 1984, J. Butze (6 USNM 
ENT 00028738 UNAM). Mexico: Teques- 
quinahuac, Cerro Tlaloc, 12 Oct 1983, M. 
Paz (3 6 USNM ENT 00028742 UNAM); 
Tequesquinahauc, Encinar, 12 Oct 1983, 
G. Sampedro (6 USNM ENT 00028746 
UNAM). Morelos: Tetela del Volcan, 8 Jul 
1978, G. Aratz (2 USNM ENT 00028748 
UNAM); 6 mi E Cuernavaca, | Sep 1974, 
G. Bohart & W. Hanson (3 6 USNM ENT 
00037921—3 USNM); Cuernavaca-Cuautla 
Highway, Canyon de Lobos, 26 Nov 1965, 
M. W. McFadden (2 USNM ENT 
00037971 USNM) COSTA RICA. Puntar- 
enas: Gravel pit near Las Alturas, 8-57-9N 
82-51-1W GPS, 1,600 m, 16 Aug 1995, M. 
A. Metz (6 USNM ENT 00021672 USNM); 
Las Alturas, Cerro Chai, 2,100 m, 14 Aug 
1995, T. Pape (6 USNM ENT 00037984 
NRS); Send. a c. Pittier, 1 km de la Estacion, 
1,800—2,000 m, LS 331800.577400, 8—20 
July 1997, M. M, Moraga Red, Lot#47388 
(2 INBIOCRIO02567298 INBIO). 

Discussion.—Associated with a pupari- 
um in the AMHH are the notes from its 
collector (Marston Bates) which declares 
the adult was reared “from aphis colony on 
Citrus (probably Aphis sp.),” the larva be- 


792 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ing a “beautiful rose and blue, looking 
much like a limacodid.”’ 

Ocyptamus willistoni is readily recog- 
nized by the lack of yellow pollinose vittae 
on the mesonotum, the red apex of the ab- 
domen, and the blackish apical wing mac- 
ula. 


ACKNOWLEDGMENTS 


We thank Drs. A. O. Bachmann and Ad- 
riana Olvia, Museo Argentino de Ciencias 
‘‘Bernardino Rivadavia,’’ Buenos Aires 
(MACN); A. Contreras-Ramos, Instituto de 
Biologia, Universidad Nacional Aut6noma 
de Mexico, Mexico (UNAM); Jeffrey M. 
Cumming and J. R. Vockeroth, Canadian 
National Collection, Agriculture Canada, 
Ottawa (CNC); David Grimaldi, American 
Museum of Natural History, New York 
(AMNH); Paul Hanson, Museo de Insectos, 
Universidad de Costa Rica, San Pedro de 
Montes de Oca, San Jose (MIUCR); Steven 
Krauth, Department of Entomology, Uni- 
versity of Wisconsin, Madison (UWEC); 
Ruth Contreras-Lichtenberg, Naturhisto- 
risches Museum Wien, Vienna (NMW); 
Thomas Pape, Naturhistoriska Riksmuseet, 
Stockholm (NRS); Philip Perkins, Museum 
of Comparative Zoology, Cambridge 
(MCZ); Richard Vane-Wright, The Natural 
History Museum, London (BMNH); E C. 
do Val, Museu de Zoologia, Universidade 
de Sao Paulo, SAo Paulo (MZUSP); for per- 
mission to study material in their care. Oth- 
er collection acronyms used are USNM for 
United States National Museum (= the Na- 
tional Museum of Natural History, Smith- 
sonian Institution), Washington, D.C. and 
INBIO for Institudo Nacional de Biodiver- 
sidad, Santo Domingo de Heredia. 

We are indebted to our colleague, Jestis 
A. Ugalde, for rearing a good portion of the 
tristani material (flies and parasites). 

We also thank Drs. Stephen D. Gaimari 
and Wayne N. Mathis, Smithsonian Insti- 
tution, Washington, D.C.; Alexander Kon- 
stantinov, Allen L. Norrbom, and Michael 
E. Schauff, Systematic Entomology Labo- 
ratory, USDA, Washington, D.C. & Belts- 


ville, Md., for their critical reviews of the 
manuscript. Finally, thanks are due to Ti- 
tana Litwak and Leonardo Donzo (figs. 20— 
23) for the fine illustrations. Titana Litwak 
as copyright holder of the color habitus 
hereby makes this image available for non- 
commerical and scientific use only. 

This work began as a review of the Me- 
soamerican species done as a class project 
by the junior author under the direction of 
the senior, who later expanded the manu- 
script to cover the subgenus as a whole. 


LITERATURE CITED 


As precise dates of publication are fre- 
quently needed to determine priority, these 
are given in brackets following the citation 
(see Thompson et al. 1999 for more de- 
tails). 


Aldrich, J. M. 1905. A catalogue of North American 
Diptera. Smithsonian Miscellaneous Collections 
46(2), 680 pp. [before 1905.05.25] 

Arnaud, P. H., Jr. and T. C. Owen. 1981. Charles How- 
ard Curran (1894-1972). Myia 2, 111 + 393 pp. 
[1981.10.23] 

Bigot, J, M. E 1883. Dipteres nouveaux ou peu con- 
nus. 22e partie, XXXII: Syrphidi (2e partie). Es- 
peces nouvelles, no. I. Annales de la Société En- 
tomologique de France (6)3: 315—356. 

Brethes, J. 1905. Insectos de Tucuman. Anales del Mu- 
seo Nacional de Buenos Aires (3)4: 329—347. 
Bruch, C. 1923. Observaciones biol6gicas acerca de 
“Salpingogaster nigriventris’ Bigot (Dipteros- 

Sirfidos). Physis 7: 1-6. 

Cooper, B. E. and J. M. Cumming. 1993. Diptera types 
in the Canadian National Collection of Insects. 
Part 2 Brachycera (exclusive of Schizophora). Re- 
search Branch, Agriculture Canada Publication 
1896/B, iii + 105 pp. 

Coquillett, D. 1910. The type-species of the North 
American genera of Diptera. Proceedings of the 
United States National Museum 37: 499-647. 
[1910.08.04] 

Curran, C. H. 1929. New Syrphidae and Tachinidae. 
Annals of the Entomological Society of America 
22: 489-510. [before 1929.10.15 receipt date 
Smithsonian Libraries] 

. 1941. New American Syrphidae. Bulletin of 
the American Museum of Natural History 78: 
243-304. [1941.08.07] 

Fluke, C. L. 1950. The male genitalia of Syrphus, Epis- 
trophe and related genera (Diptera: Syrphidae). 
Transactions of the Wisconsin Academy of Sci- 
ences, Arts and Letters 40: 115-148. 


VOLUME 102, NUMBER 4 


. 1956. Catalogue of the family Syrphidae in 
the Neotropical Region. Revista Brasileira de En- 
tomologia 6: 193-268. [1956.12.10] 

. 1957. Catalogue of the family Syrphidae in 

the Neotropical Region. Revista Brasileira de En- 

tomologia 7: 1-181. [1957.06.20] 

. 1958. A study of the male genitalia of the 
Melanostomini (Diptera: Syrphidae). Transactions 
of the Wisconsin Academy of Sciences, Arts and 
Letters (1957) 46: 261-279. 

Hanson, P. E. and I. D. Gauld. 1995. The parasitoid 
and its interactions with its host. Section 2.5, pp. 
45—67. In Hanson, P. E. and I. D. Gauld, eds., The 
Hymenoptera of Costa Rica. Oxford Univesity 
Press, xx + 893 pp. 

Hull, E M. 1943a. New species of syrphid flies in the 
National Museum. Journal of the Washington 
Academy of Sciences 33: 39—43. [1943.02.15] 

. 1943b. Two new species of Baccha (Diptera: 

Syrphidae). Proceedings of the Entomological So- 

ciety of Washington 45: 50-51. [1943.03.06] 

. 1943c. New species of Syrphidae (Diptera) of 

the genera Baccha and Mesogramma. Entomolog- 

ical News 54: 89-92. [1943.04.27] 

. 1943d. The New World species of the genus 

Baccha. Entomologica Americana 23: 42—99. 

[1943.04.30] 

. 1943e. New species of Baccha and related 

flies. Entomological News 54: 135-140. 

[1943.06.07] 

1949. The genus Baccha from the New 
World. Entomologica Americana (1947) 27: 89— 
291. [1949.02.04] 

Kertész, K. 1910. Catalogus dipterorum hucusque des- 
criptorum. Budapest, Vol. 7, 470 pp. [1910.06.??] 

Knutson, L. V. 1971. Puparia of Salpingogaster con- 
opida and S. texana, with notes on prey (Diptera: 
Syrphidae). Entomological News 82: 29-38. 
[1971.05.28] 

Loew, H. 1863. Diptera Americae septentrionalis in- 
digena. Centuria tertia. Berliner Entomologische 
Zeitschrift 7: 1-55. [1863.06.??] 

Macquart, J. 1834. Histoire naturelle des Insectes.— 
Dipteres. Diptera, Vol. 1, 578 pp., 12 pls. Jn Roret, 
N. E., ed., Collection des suites 4 Buffon. Paris 
[1834.03.31] 

Noyes, J. S. 1995. Encyrtidae. Section 11.5, pp. 300— 
309. In Hanson, P. E. and I. D. Gauld, eds. The 
Hymenoptera of Costa Rica. Oxford Univesity 
Press. xx + 893 pp. 

Sack, P. 1920. Die Gattungen Salpingogaster Schiner 
und Meromacrus Rondani. Zoologische Jahrbuch- 
er. Abteilung fur Systematik Okologie und Geo- 
graphie der Tiere 43: 235-272. 

. 1921. Dr. L. Ziircher’s Dipteren-Ausbeute aus 

Paraguay: Syrphiden. Archiv fiir Naturgeschichte 

3: 127-149. 


WES 


Schiner, J. R. 1868. Diptera. vi + 388 pp., 4 pls. In 
Wullerstorf-Urbair, B. von, in charge, Reise der 
osterreichischen Fregatte Novara. Zoologie 2(1)B. 
K. Gerold’s Sohn, Wien. [1868.04.30] 

Rotheray, G. E. 1986. Colour, shape and defence in 
aphidophagous syrphid larvae (Diptera). Zoolog- 
ical Journal, Linnaean Society 88: 201-216. 
[1986.11.12] 

Thompson, E C. 1981. The flower flies of the West 
Indies (Diptera: Syrphidae). Memoirs, Entomolog- 
ical Society of Washington 9, 200 pp. [1981.??.??] 

. 1999. A key to the genera of the flower flies 

(Diptera: Syrphidae) of the Neotropical Region in- 

cluding descriptions of new genera and species 

and a glossary of taxonomic terms. Contributions 

on Entomology, International 3: 319-378. 

[1999.08.23] 

. 2000. Nearctic flower flies (Diptera: Syrphi- 
dae). Myia 11 (In press.) 

Thompson, E C., N. L. Evenhuis, and C. W. Sabrosky. 
1999. Bibliography [of family-group names in 
Diptera]. Myia 10: 361—574. [1999.03.15] 

Thompson, FE C., J. R. Vockeroth, and Y. S. Sedman. 
1976. Family Syrphidae. /n Papavero, N., ed., A 
Catalogue of the Diptera of the Americas South 
of the United States. Departmento de Zoologia, 
Secretaria da Agricultura. Sao Paulo, Brazil. Fas- 
cicle 46, 195 pp. [1976.08.09] 

Tristan, J. F 1897. Insectos de Costa Rica. Museo Na- 
cional de Costa Rica, Tipografia Nacional, San 
José, Costa Rica, 21 pp. 

Vockeroth, J. R. 1969. A revision of the genera of the 
Syrphini (Diptera: Syrphidae). Memoirs. Ento- 
mological Society of Canada 62, 176 pp. 
[1969.07.15] 

Williston, S. W. 1886. Catalogue of the described spe- 
cies of South American Syrphidae. Transaction of 
the American Entomological Society 14: 308— 
324. 

. 1887. Synopsis of the North American Syr- 

phidae. Bulletin. United States National Museum 

(1886) 31, xxx + 335 pp. [before 1887.06.30] 

. 1891. Fam. Syrphidae, pp. 1-56. Jn Godman, 
E D. and Salvin, O., eds., Biologia Centrali-Amer- 
icana. Zoologia-Insecta-Diptera, Vol. 3, 127 pp. 
[1891.12.??] 

Wolcott, G. N. 1923. “‘Insectae Portoricensis.”’ A pre- 
limnary annotated check-list of the insects of 
Puerto Rico, with descriptions of some new spe- 
cies. Journal of the Department of Agriculture, 
Puerto Rico 7(1), 312 pp. 

. 1936. “‘Insectae Borinquenses.”’ A revised an- 

notated checklist of the insects of Puerto Rico. 

Journal of Agriculture, University of Puerto Rico 

2(1), 601 pp. 

. 1948. The insects of Puerto Rico. Diptera. 

Journal of Agriculture, University of Puerto Rico 

32(3): 417-532. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 794-801 


A NEW SPECIES OF TRIASPIS HALIDAY (HYMENOPTERA: BRACONIDAE) 
PARASITIC ON THE PEPPER WEEVIL, ANTHONOMUS EUGENI CANO 
(COLEOPTERA: CURCULIONIDAE) 


ROBERT A. WHARTON AND VICTOR LOPEZ-MARTINEZ 


(RAW) Department of Entomology,Texas A&M University, College Station, TX 77843, 
U.S.A. (e-mail: rawbaw2@acs.tamu.edu); (VL-M) Colegio de Postgraduados, IFIT-En- 
tomologia Km 35.5 carr. México-Texcoco Montecillo, edo. de México C.P. 56230 Mexico 


Abstract.—A new species of braconid wasp, Triaspis eugenii, is described. This species 
is a parasitoid of the pepper weevil, Anthonomus eugenii, in Nayarit, Mexico. It is being 
investigated as part of a classical biological control program directed against this pest in 
Florida. Diagnostic features are provided for separating this species from other species of 
Triaspis attacking weevil pests. The overall similarity to another pepper weevil parasitoid 


in the genus Urosigalphus is also noted. 


Key Words: 
control, Mexico 


The genus Triaspis was first described by 
Haliday (1835), and is currently placed in 
the tribe Brachistini of the subfamily Hel- 
coninae by most workers (Mason 1974, 
Sharkey 1997). Though confused with Si- 
galphis Latreille for nearly 80 years, the 
true identity of Triaspis was fixed in the 
early 1900s and has remained stable ever 
since (Martin 1955). Triaspis is character- 
ized by the presence of a completely fused 
carapace and the absence of the r-m cross- 
vein in the fore wing. In typical Triaspis, 
the carapace retains the two transverse fur- 
rows: all that remain of the sutures that 
originally separated the three segments 
comprising the carapace. Schizoprymnus 
Foerster, sometimes recognized as a sepa- 
rate genus and sometimes treated as a sub- 
genus of Triaspis, is separated from typical 
Triaspis by the absence of the two trans- 
verse furrows, at least dorsally. Most other 
brachistines have at least the first two me- 
tasomal segments separated by a flexible 
suture. The only exception in the New 


Braconidae, Helconinae, Brachistini, pepper weevil, parasitoid, biological 


World is the genus Urosigalphus Ashmead, 
otherwise distinguished by having the outer 
tarsal claw of the hind leg much larger than 
the inner claw. In the New World, various 
species of Triaspis have been reared from 
the same weevil hosts as other brachistines 
in the genera Urosigalphus, Nealiolus Ma- 
son, and/or Aliolus Say. 

The most recent revisions or reviews of 
Triaspis include those of Martin (1955) for 
North America, Papp (1984) for Australia, 
Tobias (1986) for the European part of the 
former USSR, Priore and Tremblay (1987) 
for Italy, Chou and Hsu (1996) for Taiwan, 
and Belokobylskij (1998) for Eastern Rus- 
sia. The neotropical fauna has never been 
revised and isolated descriptions of only 
five species have been published to date 
(Shenefelt 1970). A Palaearctic species at- 
tacking the pea weevil, Bruchus pisorum 
(L.), has also been introduced to the Neo- 
tropics (Shenefelt 1970). 

In general, members of the genus Trias- 
pis are poorly known biologically (Shaw 


VOLUME 102, NUMBER 4 


and Huddleston 1991). Nevertheless, a few 
species have been reared from weevils, bru- 
chids, and anthribids, and those that are best 
studied oviposit in the host egg and emerge 
from late instar larvae. The species de- 
scribed here has been reported as the most 
abundant parasitoid of the pepper weevil, 
Anthonomus eugenii Cano, in the Mexican 
state of Nayarit, attaining 50% parasitism 
in some of the samples from commercial 
hot pepper crops there (Mariscal et al. 
1998). The species is currently being tested 
for possible use in the biological control of 
the pepper weevil in Florida. Information 
on the host weevil has recently been sum- 
marized by Clark and Burke (1996). 


MATERIAL AND METHODS 


All of the material used for this study 
was collected in Nayarit, Mexico, and was 
reared from peppers containing the pest 
weevil A. eugenii. The majority of the spec- 
imens were collected by Eugenio Mariscal 
and by Marco Toapanta and Phil Stansly as 
part of a biological control program through 
the University of Florida, in cooperation 
with Mario Urias of the National Institute 
of Agriculture and Forestry Research (IN- 
IFAP). 

Terminology for the desciption generally 
follows Sharkey and Wharton (1997). Ad- 
ditionally, the 3 segments of the carapace 
are referred to as Tl, T2, and T3. Total 
length of ovipositor was determined by dis- 
section of 10 individuals, and comparing 
the total length with that of the dissected 
metasoma. Approximate length of oviposi- 
tor is also indicated in the description, as 
distance protruding beyond apex of cara- 
pace. The carapace is bordered by a thin 
flange called the marginal lamella (Martin 
1955). Variation in quantitative traits is in- 
dicated by a range and mean, based on 10 
males and 10 females. 

Figures were produced using a scanning 
electron microscope (SEM). All SEM im- 
ages were shot at 10kv on a JOEL JSM- 
5600. Specimens were coated with about 20 


795 


nm gold, using a Hummer 6.2 sputtering 
unit with argon as a source gas. 


RESULTS 


Triaspis eugenii Wharton and 
L6épez-Martinez, new species 
(Figs. 1-17) 


Holotype female.—Top label = ““MEX- 
ICO: Nayarit Mpio. Santiago Ixcuintla, 
Puerta de Mango, 24.11.1999 Toapanta & 
Stansly”’ Second label = “reared from An- 
thonomus eugenii on Serrano pepper 
emerged 7.iv.1999°’. Deposited in collec- 
tion of Universidad Nacional Aut6noma de 
Mexico, Mexico City. Paratypes: 9 females, 
10 males, same data as holotype. 25 fe- 
males, 26 males, ““MEXICO: Nayarit Bal- 
adero 2.11.1997 E. Mariscal’’; 3 females, 
“Santiago Ixc. Nay. 23 febrero 1997 E. 
Mariscal Chile Serrano 11 msnm Hosp. An- 
thonomus eugenii’’. Paratypes deposited in 
the entomological collections of the follow- 
ing institutions: Texas A&M University, 
College Station; The Natural History Mu- 
seum, London; Universidad Aut6énoma de 
Yucatan, Mérida; Universidad Postgradua- 
dos, Montecillos; Universidad Aut6noma 
de Nuevo Leon, Monterrey; The Canadian 
National Collection, Ottawa; El Colegio de 
la Frontera Sur, Tapachula; National Mu- 
seum of Natural History, Smithsonian In- 
stitution, Washington, DC; and Instituto de 
Ecologia, Xalapa. 

Diagnosis.—This species is characterized 
by the nearly bare mesoscutum and scutel- 
lum (Fig. 13), the propodeum with a very 
short anterior face and sharply declivous 
posterior face (Figs. 10—11), the pattern of 
sculpture on the metasoma (Figs. 14-17), 
and a clypeus that is twice wider than high 
(Figs. 1-2). Triaspis eugenii is most similar 
to Triaspis azteca Martin from Mexico and 
especially Triaspis vestiticida Viereck from 
Peru. The flagellum is uniformly dark in 7. 
azteca but the basal flagellomeres are pale 
(especially ventrally) in 7. eugenii and most 
specimens of T. vestiticida. The mesonotum 
is almost uniformly sparsely setose in T. 


796 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


se 


} 


3G tira 5 
Figs. 1-6. Head of Triaspis eugenii. 1, Clypeus in frontal view, arrows showing median and lateral lobes. 
2, Face in frontal view showing setal pattern. 3, Anterior-dorsal view of clypeus showing rugulose and pitted 


surface. 4, Head in dorsal view, arrow showing depression on frons immediately posteriorad antennal base. 5, 
Mandible. 6, Lateral view of head. 


9) 


vestiticida but largely bare in T. eugenii. Of giniensis, the species described here differs 
the North American species treated by Mar- in having more extensive sculpture on the 
tin (1955), 7. eugenii most closely resem- carapace, and a taller, narrower clypeus 
bles Triaspis virginiensis Ashmead and (clypeus ‘“‘transverse’”’ in T. virginiensis, 
Triaspis rectangulata Martin. From T. vir- about 0.75 X width of face). From T. rec- 


VOLUME 102, NUMBER 4 


ee 
S36 bre 


Figs. 7-9. 


197) 


Antenna of female Triaspis eugenii. 7, Composite view of flagellum (scape and pedicel absent). 


8, Basal 3 flagellomeres showing relative lengths of the 3 segments and patch of short sensilla (arrow) on 
flagellomere 1. 9, Flagellomeres 7—9, showing patch of irregular setae and placode sensilla; see Fig. 7 for scale. 


tangulata, it differs in antennal coloration 
and shape of the head. 

Description.—Female. Length: 1.85—2.15 
mm. 

Head: Polished; somewhat trapezoidal, 
slightly narrowed ventrally in frontal view 
(gena thus not inflated: Fig. 2); vertex, oc- 
ciput, temple, and gena sparsely setose 
(Figs. 4, 6), the setae arising from indistinct 
punctures; face smooth, more densely cov- 
ered with ventrally and ventral-medially di- 
rected setae (Fig. 2); frons (Fig. 4) sparsely 
setose laterally, bare medially, with shallow 
depressions posteriorad scape on either side 
of a low, median ridge, depressions weakly 
sculptured in part, frons otherwise polished; 
eye large, about twice longer than temple 
in dorsal view, with temple strongly reced- 
ing behind eye; malar space 0.25—0.3 X eye 
height, 0.4—0.45 xX height of face; face 
1.65—1.85 (m = 1.75) X wider than high; 


epistomal suture deeply impressed through- 
out; clypeus narrow: width 0.57—0.65 (m = 
0.62) X width of face, about twice wider 
than high, surface uneven and distinctly 
punctate (Fig. 3), ventral margin sinuate: 
with weak, broadly rounded median lobe 
and a large, very obtuse angulation near the 
base of each mandible (Figs. 1, 5), weakly 
convex in profile; mandible (Fig. 5) lacking 
obvious sculpture; ocelli not surrounded by 
an impressed groove; antenna (Fig. 7) with 
18-19 segments; first two flagellomeres 
equal in length (Fig. 8), flagellomeres 3—9 
gradually shortening towards apex, flagel- 
lomeres 9 through 14—15 bead-like (Fig. 9), 
nearly as wide as long; maxillary palps 
short, about equal in length to eye height, 
6-segmented; labial palps with only three 
apparent segments. 

Mesosoma: Pronotum dorsally deeply 
pitted along posterior margin, the sculpture 


798 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 10-15. 


11, Lateral view of thorax and propodeum. 12—13, Dorsal view of mesosoma. 14—15, Dorsal view of metasoma 
showing slightly different degrees of development of sculpture. 


weakening laterally, extending for a vary- 
ing distance ventrally as a crenulate line; 
lateral face of pronotum (Fig. 11) bare me- 
dially, largely unsculptured in most speci- 
mens except at extreme ventral and poste- 
rior corners. Mesoscutum with anterior de- 
clivity and extreme base of notauli setose 
(Fig. 11), with 1—2 rows of scattered setae 
along notauli and around lateral margin of 
scutum (Figs. 12—13), median and lateral 
lobes otherwise bare and highly polished; 
notauli very finely crenulate and weakly 


Triaspis eugenii. 10, Lateral view of body, showing relative lengths of antenna and ovipositor. 


impressed throughout, very narrowly sepa- 
rated posteriorly, the area of convergence 
completely impunctate (Fig. 13); scutellar 
sulcus finely rugulose and with a distinct 
midridge; scutellum polished, nearly bare; 
mesopleuron (Fig. 11) bare medially, setose 
only along anterior and posterior margins, 
dorsally along subalar depression, and ven- 
trally beneath sternaulus; sternaulus shal- 
low, sinuate, unsculptured or nearly so. Me- 
tanotum (Figs. 11, 13) short, length along 
midline, when viewed in profile (Fig. 11), 


VOLUME 102, NUMBER 4 


Figs. 16-17. 
16, Posterior-lateral view. 17, Posterior view. 


equal to or shorter than median carina of 
propodeum; metapleuron densely and uni- 
formly rugose beneath a mat of short, white 
setae. Propodeum (Figs. 10—12) very short 
anteriorly, steeply declivous posteriorly; 
short-setose throughout, the sculpture read- 
ily visible through the setae; with well-de- 
fined pentagonal areola occupying roughly 
median third of declivity, declivity (includ- 
ing areola) otherwise irregularly carinate- 
rugulose, the sculpture varying in intensity, 
posterior-lateral ridge distinctly protruding. 
Hind femur short, broad, 2.85—3.1 (m = 
3.0) X longer than wide; tarsal claw with 
distinct basal tooth. Fore wing 2.4—2.5 xX 
longer than maximal width; basal cell 
sparsely setose; 1M distinctly bowed; Rla 
1.55—2.1 (m = 1.8) X longer than R1b, the 
latter not extending to wing tip; m-cu 1.7— 
2.3 (m = 1.9) X longer than (RS+M)b; 
lcu-a distinctly postfurcal, 1.15—2.5 (m = 
1.5) X longer than 1CUa; 2cu-a absent, 2- 
1A usually very short, thus Ist subdiscal 
cell broadly open posterior-distally. 
Metasoma: Approximately equal in 
length to mesosoma. Carapace short and 
broadly oval, in dorsal view 1.25—1.35 (m 
= 1.27) X longer than maximum width; ex- 
tensively sculptured; Tl and T2 carinate 
over a finely rugulose and variously pitted 
background, the surface of T1+T2 usually 
(80%) completely sculptured (Figs. 15—16), 
more rarely with sculpture evanescent pos- 


799 


‘ a Se 
EG nt ee ae 


afl wit : 


Metasoma of Triaspis eugenii showing marginal lamella and apically sinuate carapace margin. 


terior-medially adjacent T2/T3 suture (Fig. 
14), sculpture usually more strongly cari- 
nate laterally, with at least some carinae ex- 
tending across the foveolate T1/T2 suture; 
T3 polished and unsculptured over medial 
0.2—0.4 (Fig. 17), strigose laterally, T2/T3 
suture (Figs. 14—15) complete, very broadly 
u-shaped, finely crenulate; dorsal carinae of 
T1 distinctly elevated above surrounding 
sculpture and weakly converging over basal 
half, separated by about 0.4 X carapace 
width at this point, parallel-sided and grad- 
ually weakening over distal half, usually 
reaching suture as a pair of simple carinae 
similar in appearance to adjacent carinate 
sculpture; carapace somewhat truncate pos- 
teriorly, with distinct concavity mid-ven- 
trally (Fig. 17); marginal lamella (Figs. 10, 
16—17) present as a very narrow ridge along 
T1 and T2, widening posteriorly on T3, ex- 
tending completely around posterior margin 
of carapace as a broad, thin flange. Ovipos- 
itor 1.15—1.4 (m = 1.25) X longer than me- 
tasoma; at rest, extending roughly 0.7 X 
carapace length beyond apex (Fig. 14). 
Color: Black; tegula mostly black except 
outer margin brown; antenna dark brown 
with scape, pedicel, and basal 3—6 flagel- 
lomeres yellowish ventrally; clypeus vari- 
ously dark reddish brown; mandible yellow 
except apically; palps pale yellow to whit- 
ish; legs patterned as follows: hind coxa 
dark brown to black basally and posteriorly, 


800 


yellow anteriorly and apically, fore and mid 
coxae more extensively yellow; trochanters 
yellow; hind femur yellow with broad, sub- 
apical brown band, band somewhat more 
diffuse and largely limited to dorsal and 
posterior sides of mid femur, fore femur 
yellow with dark brown spot dorsally over 
much of basal half; hind tibia gradually 
darkening from base to apex, at least basal 
fourth dark brown, fore and mid tibiae yel- 
low; tarsi brown to light brown dorsally, 
yellowish ventrally. 

Male.—As in female except face 1.75— 
2.0 (m = 1.85) X wider than high; antenna 
with 19-20 segments, flagellomeres grad- 
ually becoming shorter towards apex, but 
subapical flagellomeres longer and usually 
not as distinctly bead-like as in female, first 
flagellomere shorter, 0.75—0.9 (m = 0.8) X 
length of second. Hind femur somewhat 
more slender, 2.95—3.55 (m = 3.25) X lon- 
ger than maximum width. Carapace dis- 
tinctly narrower, in dorsal view 1.3—1.45 (m 
= 1.4) X longer than maximum width. Sub- 
apical brown band on hind femur often 
more diffuse, fore and mid femora more ex- 
tensively yellow. 

Discussion.—Martin (1955), in his revi- 
sion of the North American species, delin- 
eated several species groups in Triaspis. 
The species described here, however, is not 
readily assignable to any of Martin’s infor- 
mal groups. To accommodate the neotrop- 
ical species in general, species groups will 
have to be re-defined using a larger suite of 
characters including the propodeal structure 
and shape of the clypeus. The sculpture of 
the carapace is of value in separating some 
species and species groups, but examination 
of the large amount of reared material avail- 
able to us indicates that caution is needed 
when using sculpture alone to identify iso- 
lated individuals. In T. eugenii, for exam- 
ple, the sculpture underlying the carinations 
may vary from rugulose and distinctly 
punctate to nearly smooth. 

The two previously described neotropical 
species that most closely resemble T. eu- 
genii have both been reared. Triaspis azteca 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


attacks the bean pod weevil, Apion god- 
mani Wagner (Martin 1952, Perez 1985), 
and the description of 7. vestiticida was 
based on material reared from Anthonomus 
vestitus Boheman infesting cotton (Viereck 
1912). 

Members of the genus Triaspis are su- 
perficially similar to members of the genus 
Urosigalpus, since both have a carapace- 
like abdomen and similar wing venation. 
Since both can be reared from the same 
weevil hosts (including pepper weevil), it is 
important to exercise care when identifying 
reared material. As noted above, the species 
of Urosigalphus have an unusually large 
outer claw on the hind leg. 

Etymology.—tThe species epithet is taken 
from the only known host, Anthonomus eu- 
genil. 


ACKNOWLEDGMENTS 


We are grateful to Ed Riley and Horace 
Burke (Texas A&M University) for infor- 
mation on weevil hosts of 7. azteca, Terry 
Junek for assistance with the literature, and 
Dave Smith (USDA/ARS, Systematic En- 
tomology Laboratory) for faciliting exami- 
nation of the Triaspis collection at the Na- 
tional Museum of Natural History. Phil 
Stansly, Marco Toapanta, Eugenio Maris- 
cal, José Antonio Sanchez-Garcia, and Jor- 
ge Leyva-Vazquez kindly made material 
available for the description, and provided 
additional information on hosts and the bi- 
ological control program. Special thanks to 
Matt Buffington (Texas A&M University) 
and Jim Ehrman for preparation of the im- 
ages. All SEM work was done by Jim Ehr- 
man at the Digital Microscopy Facility, 
Mount Allison University, Sackville, NB, 
Canada. This work was supported in part 
by the Texas Agricultural Experiment Sta- 
tion. The Texas Parks and Wildlife Depart- 
ment is gratefully acknowledged for per- 
mission to collect in state parks (permits 
39-49, 41-95, and 10-98), from which we 
obtained specimens of Triaspis useful for 
comparison of species-level characteristics. 


VOLUME 102, NUMBER 4 


LITERATURE CITED 


Belokobylskij, S. A. 1998. 12 Subfamily Brachistinae 
(Calyptinae), pp. 440—489. Jn Ler, P. A., ed., Key 
to the Insects of Russian Far East, Vol. ITV. Neu- 
ropteroidea, Mecoptera, Hymenoptera. Pt. 3. 
Vladivostok: Dal’nauka, 708 pp. [In Russian] 

Chou, L.-y. and T.-C. Hsu. 1996. The Braconidae (Hy- 
menoptera) of Taiwan. 7. Subtribe Triaspina. Jour- 
nal of Agricultural Research of China 45: 436— 
497. 

Clark, W. E. and H. R. Burke. 1996. The species of 
Anthonomus Germar (Coleoptera: Curculionidae) 
associated with plants in the family Solanaceae. 
Southwestern Entomologist Supplement 19: 1— 
114. 

Haliday, A. H. 1835. Essay on parasitic Hymenoptera 
of the Ichneumones Adsciti (cont.). Entomological 
Magazine 3: 121-147. 

Mariscal Mejorado, E., J. L. Leyva Vazquez, and R. 
Bujanos Muniz. 1998. Parasitoides del picudo del 
chile, Anthonomus eugenii Cano (Coleoptera: 
Curculionidae), en Nayarit, México. Vedalia 5: 
39-46. 

Martin, J. C. 1952. A new braconid from Mexico. The 
Canadian Entomologist 84: 30-31. 

. 1955. A taxonomic revision of the triaspidine 
braconid wasps of Nearctic America (Hymenop- 
tera). Canada Department of Agriculture, Science 
Service, Entomology Division. 157 pp. 

Mason, W. R. M. 1974. A generic synopsis of Brach- 
istini (Hymenoptera: Braconidae) and recognition 
of the name Charmon Haliday. Proceedings of the 
Entomological Society of Washington 76: 235— 
246. 

Papp, J. 1984. First survey of the Triaspidini species 
of the Indo-Australian region (Hymenoptera: Bra- 
conidae, Calyptinae) 1. The genus Triaspis Hali- 
day. Acta Zoologica Academiae Scientarum Hun- 
garicae 30: 137-158. 


801 


Pérez, G. 1985. Himenopteros parasitoides de Apion 
spp. (Coleoptera: Curculionoidea: Apionidae) en 
Tepoztlan, Morelos. Folia Entomologica Mexi- 
cana 63: 39—46. 

Priore, R. and E. Tremblay. 1987. Contributo alla re- 
visione delle specie italiane dei generi Triaspis 
Haliday e Schizoprymnus Foerster (Hymenoptera 
Braconidae). Bollettino del Laboratorio di Ento- 
mologia Agraria ‘‘Filippo Silvestri” di Portici 44: 
47-61. 

Sharkey, M. J. 1997. Subfamily Helconinae, pp. 260— 
272. In Wharton, R. A., P. M. Marsh, and M. J. 
Sharkey, eds., Manual of the New World Genera 
of the Family Braconidae (Hymenoptera). Inter- 
national Society of Hymenopterists. Special pub- 
lication 1, 439 pp. 

Sharkey, M. J. and R. A. Wharton. 1997. Morphology 
and Terminology, pp. 19-37. In Wharton, R. A., 
P. M. Marsh, and M. J. Sharkey, eds., Manual of 
the New World Genera of the Family Braconidae 
(Hymenoptera). International Society of Hymen- 
opterists. Special publication 1, 439 pp. 

Shaw, M. R. and T. Huddleston. 1991. Classification 
and biology of braconid wasps. Handbooks for the 
Identification of British Insects 7(11): 1-126. 

Shenefelt, R. D. 1970. Braconidae 2: Heliconinae [sic], 
Calyptinae, Mimagathidinae, Triaspinae. /n Ferri- 
ére, Ch. and J. Van der Vecht, eds., Hymenopter- 
orum Catalogus (nov. ed.) 5: 177-306. 

Tobias, V. I. 1986. 7. Subfam. Brachistinae (Calypti- 
nae). Handbook to the Insects of the European 
Part of the USSR, 3, Hymenoptera, Part 4: 158— 
180. Akademia Nauk, Leningrad, 501 pp. [In Rus- 
sian] 

Viereck, H. L. 1912. Contributions to our knowledge 
of bees and ichneumon-flies, including the de- 
scription of twenty-one new genera and fifty-sev- 
en new species of ichneumon-flies. Proceedings of 
the United States National Museum 42: 613-648. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 802-815 


NEW HOST PLANT AND DISTRIBUTION RECORDS IN MEXICO FOR 
ANASTREPHA SPP., TOXOTRYPANA CURVICAUDA GERSTACKER, 
RHAGOLETIS ZOQUI BUSH, RHAGOLETIS SP., AND HEXACHAETA SP. 
(DIPTERA: TEPHRITIDAE) 


MARTIN ALUJA, JAIME PINERO, MAURILIO LOPEZ, CESAR RUIZ, ALBERTO ZUNIGA, 
ENRIQUE PIEDRA, FRANCISCO DIAZ-FLEISCHER, AND JOHN SIVINSKI 


(MA, JP, ML, CR, AZ, EP) Instituto de Ecologia, A.C., Apartado Postal 63, C.P. 91000 
Xalapa, Veracruz, México (MA, e-mail: alujam @ecologia.edu.mx); (FD-F) Departamento 
de Desarrollo de Métodos, Campania Nacional Contra las Moscas de la Fruta, 2a Avenida 
Sur No. 5 (Altos), C.P. 30700 Tapachula, Chiapas, México; (JS) Center for Medical, 
Agricultural and Veterinary Entomology, USDA ARS, 1700 SW 23rd Drive, P.O. Box 
14565, Gainesville, FL 32604, U.S.A. 


Abstract.—We report the results of a nine-year study (1990-1998), aimed at determin- 
ing the distribution of fruit flies (Diptera: Tephritidae) and identify their host plants in 
Veracruz, Mexico. Twenty Anastrepha species, Toxotrypana curvicauda and several un- 
identified Hexachaeta species were captured in McPhail traps. Among the 20 Anastrepha 
species, A. limae, A. dentata, and A. canalis are new records for Veracruz. Out of a total 
of 51 species of native, wild plants and exotic, cultivated plants collected in Veracruz, 
Mexico (representing 3,736 kg of fruit), 34 were infested by 13 Anastrepha species, 4 
plants were infested by 4 Rhagoletis species, 2 plants were infested by Toxotrypana 
curvicauda, and one by an undescribed Hexachaeta species. Our observations include 
new host plant records for Anastrepha hamata (larvae feed only on seeds), A. bahiensis, 
A. chiclayae, A. fraterculus, A. obliqua, T. curvicauda, Rhagoletis zoqui, R. sp. and Hex- 
achaeta sp. We present host plant data based on size of fruit and degree of infestation, 
provide information on local names and fruiting phenology, and discuss our findings in 
light of their practical implications and with respect to the zoogeography of Mexican fruit 
flies. 


Resumen.—Reportamos los resultados de un estudio de nueve anos (1990-1998), en- 
caminado a determinar la distribucidn de moscas de la fruta (Diptera: Tephritidae) e 
identificar sus plantas hospederas en Veracruz, México. Veinte especies de Anastrepha, 
Toxotrypana curvicauda y varias especies no identificadas de Hexachaeta fueron captu- 
radas en trampas McPhail. De estas especies A. limae, A. dentata y A. canalis representan 
nuevos reportes para Veracruz. De un total de 51 especies de plantas nativas (silvestres) 
y exOticas (cultivadas) colectadas en Veracruz, México (representando 3,736 kg de fruta), 
34 fueron infestadas por 13 especies de Anastrepha, otras 4 por 4 especies de Rhagoletis, 
2 por Toxotrypana curvicauda y una por una especie no descrita de Hexachaeta. Nuestras 
observaciones incluyen nuevas plantas hospederas para Anastrepha hamata (las larvas se 
alimentan sdlo de semillas), A. bahiensis, A. chiclayae, A. fraterculus, A. obliqua, T. 
curvicauda, Rhagoletis zoqui, R. sp. y Hexachaeta sp. Presentamos informacion sobre 
plantas hospederas basada en el tamafio de fruto y grado de infestacidén, proporcionamos 
informacion acerca de nombres locales de los frutos, su fenologia de fructificacién y 


VOLUME 102, NUMBER 4 


803 


discutimos nuestros descubrimientos en relaci6n a sus implicaciones practicas y con res- 
pecto a la zoogeografia de las moscas de la fruta presentes en México. 


Key Words: 
zoogeography 


Flies in the genera Anastrepha Schiner, 
Rhagoletis Loew, Toxotrypana Gerstacker, 
and Hexachaeta Loew are widely distrib- 
uted in the Neotropics (Norrbom et al. 
1999). In Anastrepha, there are 197 de- 
scribed species distributed from the south- 
ern USA to northern Argentina (Aluja 
1994, Norrbom et al. 2000). The host plants 
of Anastrepha were recently reviewed by 
Norrbom (2000) and Norrbom et al. (2000). 
These combined reports indicate that plant 
species in more than 75 families are used 
as hosts. For Toxotrypana curvicauda Ger- 
stacker, host plants are restricted to two 
families only: Caricaceae and Asclepiada- 
ceae. They include species such as Carica 
papaya L., Carica cauliflora Jacq., Morren- 
ia odorata Lindl., and Gonolobus sorodius 
Gray. (Mason 1922, Castrej6n-Ayala and 
Camino-Lavin 1991, Landolt 1994). For 
flies in the genus Hexachaeta, almost no 
host-plant data are available. This genus in- 
cludes 25 described species and many un- 
described species, and the few host plant 
reports available are predominantly plants 
in the family Moraceae (Norrbom et al. 
1999). 

Here, we report the results of a long-term 
study (nine years of year-round fruit collec- 
tions and trapping activities) aimed at de- 
termining the distribution of the fruit fly 
genera Anastrepha, Rhagoletis, Toxotry- 
pana, and Hexachaeta and identifying their 
host plants in the State of Veracruz, Mexi- 
co. 


MATERIALS AND METHODS 


Study sites.—During nine years (1990-— 
1998) we collected all wild or cultivated 
fruit (potential fruit fly hosts) we could find 
in 42 sites located in the state of Veracruz, 


Anastrepha, Rhagoletis, Toxotrypana, Hexachaeta, distribution, host plants, 


Mexico. Two of these 42 sites are located 
in northern Veracruz (Morgadal and Ala- 
mo), one site is located in southern Vera- 
cruz (Los Tuxtlas), and the rest (39 sites) 
are located in central Veracruz. All study 
sites, their exact location (latitude, longi- 
tude) and altitude are described in Table 1. 

Fruit sampling methods.—Ripe, and on 
occasion unripe, fruits of the plants listed 
in Table 2, were collected in areas with un- 
perturbed and perturbed native vegetation, 
in backyard gardens, and in semicommer- 
cial and commercial orchards as described 
by Aluja et al. (1987). Samples were 
placed in plastic washbowls and transport- 
ed to our laboratory in Xalapa, Veracruz, 
Mexico. There, samples were individually 
weighed and placed in plastic trays which 
in turn were placed over a plastic wash- 
bowl containing vermiculite as a pupation 
medium. 

Processing of larvae and pupae.—We in- 
spected, on a daily basis, the pupation me- 
dium in the washbowls described above and 
collected all pupae and larvae. These were 
transferred to labeled, 500 ml plastic con- 
tainers with vermiculite. All containers (one 
per sample) were kept in the laboratory at 
26 + 2°C and 60-70% RH until all adult 
flies had emerged from the pupae. 

Trapping of adult flies —Traps to survey 
adult flies were placed in Apazapan, Llano 
Grande, Monte Blanco, and Martinez de la 
Torre, Veracruz. With the exception of Mar- 
tinez de la Torre, we worked in commercial 
mango orchards surrounded by patches of 
native vegetation, coffee plantations, and 
other orchards. In Martinez de la Torre we 
worked in a commercial grapefruit (Citrus 
paradisi Macfad.) orchard. At all sites we 
placed 16 McPhail traps baited with 250 ml 


804 


of a mixture of hydrolyzed protein, water, 
and borax in each orchard. In Apazapan we 
also placed 16 traps in a commercial chi- 
cozapote (Manilkara zapota [L.] P. Royen) 
orchard adjacent to the mango orchard. Bait 
preparation, and placement and service of 
traps are described in detail in Aluja et al. 
(1996). 

Identification of adult flies and plants.— 
All flies emerging from puparia or caught 
in McPhail traps were identified at the In- 
stituto de Ecologia, A. C., by Vicente Her- 
nandez-Ortiz. Plants were dried, pressed, 
and identified by Carlos Duran, staff mem- 
ber of the IXAL herbarium at the Instituto 
de Ecologia, A. C., in Xalapa, Veracruz. 
Voucher specimens of plants and insects not 
already represented in the permanent col- 
lections were kept in the IXAL herbarium 
and IXAL permanent insect collection. All 
plant names used here correspond to those 
listed in the USDA’s GRIN online database 
(USDA 2000). 


RESULTS 


Fruit sampling.—A total of 3,736 kg of 
fruit, representing 51 species from 22 plant 
families, was processed during this study 
(Table 2). Of these, 42 species from 17 fam- 
ilies were found to be natural hosts of 13 
Anastrepha species, 4 Rhagoletis species, 
Toxotrypana curvicauda, and an undescri- 
bed Hexachaeta species (Table 3). The 
fruiting phenology of all of these plant spe- 
cies is described in Table 4. 

The more polyphagous species were A. 
ludens (Loew) (8 plant species attacked), 
followed by A. obliqua (Macquart) and A. 
fraterculus (Wiedemann) with 7 plant spe- 
cies attacked. In Apazapan we found Myr- 
ciaria floribunda (H. West ex Will.) O. 
Berg (Myrtaceae) simultaneously infested 
by three species of Anastrepha: A. frater- 
culus, A. obliqua, and A. bahiensis Lima 
(new host plant records for the latter two 
species). We were also able to confirm that 
Terminalia catappa L. (Combretaceae) is a 
natural (i.e., field-infested) host plant of A. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


fraterculus (first report in the State of Ve- 
racruZz). 

In contrast to the polyphagous A. obli- 
qua, A. ludens and A. fraterculus, A. aphel- 
ocentema Stone and A. alveata Stone were 
monophagous species attacking Pouteria 
hypoglauca (Standley) Baehni (Sapotaceae) 
and Ximenia americana L. (Olacaceae), re- 
spectively. Anastrepha hamata (Loew) was 
an oligophagous species attacking seeds of 
both Chrysophyllum mexicanum Brandegee 
ex. Stand. and Pouteria campechiana 
(Kunth) Baehni fruits, belonging to the 
family Sapotaceae. Rhagoletis zoqui Bush, 
R. pomonella Walsh, R. turpiniae Hernan- 
dez and R. sp. adults were obtained from 
Juglans pyriformis Liebm., Crataegus mex- 
icana DC., Turpinia insignis (H.B. & K.) 
Tul., and Solanum ionidium Bitter, respec- 
tively. Toxotrypana curvicauda adults were 
obtained from Gonolobus niger (Cav.) R. 
Br. (Asclepiadaceae) fruits collected in the 
Llano Grande region. Hexachaeta sp. adults 
attacked Trophis mexicana (Liebm.) Bur. 
(Moraceae) fruits in the Monte Blanco re- 
gion (details of all the above are in Table 
5). 

Infestation patterns (i.e., number of lar- 
vae per kg of fruit) are described in Table 
5. Notably, the smaller, wild fruits were the 
most infested (Table 5, Fig. 1). 

The highest infestation levels (larvae/kg 
of fruit) were found in Spondias mombin 
L., followed by Ximenia americana L. 
(fruits in the families Anacardiaceae and 
Olacaceae attacked by A. obliqua and A. al- 
veata, respectively), while the lowest infes- 
tation levels were found in Jnga jinicuil G. 
Don (Leguminoseae), infested by A. dis- 
tincta Greene, followed by Tabernaemon- 
tana alba Mill. (Apocynaceae) attacked by 
A. cordata Aldrich (Table 5). 

Trapping.—We captured 20 Anastrepha 
species in McPhail traps. Of these, 16 spe- 
cies were captured in Apazapan, 12 species 
in Llano Grande, 14 in Monte Blanco and 
11 in Martinez de la Torre (Table 6). In the 
mango orchards, A. obliqua was by far the 
most abundant species, followed by A. lu- 


VOLUME 102, NUMBER 4 805 
Table 1. Study site locations. 
Northern Western Altitude 
No. Study site latitude longitude (meters) 
1 El Conchal 19°04’ 96°06’ 17 
D La Mancha 19°35’ 96°22’ 18 
3 Cardel 193227 96°22’ 31 
4 Playa Azul 19D 96°23’ 47 
5 Playa Oriente LOPE 96°19’ 121 
6 Paso de Ovejas 19°17’ 96°26’ 122 
oT Morgadal (Poza Rica) 20°24’ 97°21’ 128 
8 Alamo OSS) 97°40’ 180 
9 Puente Nacional 19°19’ 96°28’ 198 
10 Los Tuxtlas 18°25’ 95°06’ 200 
11 Coyolar Ome 96°36’ 228 
12 El Aguaje 1925! 96°36’ 250 
13 El Crucero Om We 96°35’ 281 
14 Actopan 19°30’ 96°36’ Sui 
i) Plan del Rio 1hOF23% 96°39’ 319 
16 Rinconada 1S 2 96°33’ B22, 
17 Apazapan 19°19’ 96°42’ 331 
18 Martinez de la Torre 19°58’ 96°47’ 400 
19 Emiliano Zapata eal 96°33’ 416 
20 Cerro Gordo 19"25¢ 96°41’ 559 
ay Llano Grande 19°22’ 96°53’ 680 
22 Corral Falso 19°27’ 96°45’ 744 
23 San Pedro 19°24’ 96°52’ 904 
24 Tejeria 19°22’ 96°56’ 924 
25 Miradores 19°28’ 96°46’ 966 
26 Limones 19°20’ 96°55’ 1,021 
DF Monte Blanco 19°23’ 96°56’ 1,026 
DS Bella Esperanza 19°26’ 96°52’ 1,057 
29 Vaqueria 19°19" 96°57’ 1,103 
30 Coatepec 19°26’ 96°57’ 1,142 
Sil Xico 19°25’ 97°00’ EAS) 
32 Teocelo 19°24’ 96°58’ 1,163 
33 Jilotepec 19°37 96°56’ 1,292 
34 Cosautlan 19°20’ 96°59’ 1,318 
35 Texin 19°20’ 97°01’ 1,464 
36 Xalapa LOSSY 96°54’ 1,468 
37] Amatla 19°19’ 97°03’ 1,478 
38 El Diez 9 2ie 97°01’ 1,498 
39 La Perla 19°19’ 97°03’ 1,578 
40 El Arenal 19°19’ 97°05’ 1,649 
41 Acajete 19°34’ 96°59’ 1,875 
42 La Joya esr 97°02’ 2,004 


dens (Monte Blanco and Llano Grande) and 
A. serpentina (Wiedemann) (Apazapan). At 
Martinez de la Torre (citrus orchard), A. lu- 
dens was the most common Anastrepha 
species captured. All of these species are 
considered agricultural pests of economic 
importance in Mexico. 

Many Anastrepha species of non-eco- 


nomic importance were also captured at all 
four trapping sites (e.g., A. bicolor [Stone], 
and A. spatulata Stone). In contrast, A. bah- 
iensis, A. dentata (Stone), A. robusta 
Greene, and A. zuelaniae Stone were found 
only in Apazapan, while A. canalis Stone 
was captured only in Monte Blanco. Adults 
of Hexachaeta spp. were commonly cap- 


806 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 2. Plant species sampled in central Veracruz, Mexico, to determine their host status to the local tephritid 


fruit flies. 


Plant family 


Scientific name 


Local common name 


Locality code* 


Anacardiaceae 


Annonaceae 
Apocynaceae 
Asclepiadaceae 
Bombacaceae 


Caricaceae 
Combretaceae 
Euphorbiaceae 
Guttiferae 
Juglandaceae 
Leguminoseae 


Malpighiaceae 
Moraceae 


Myrsinaceae 
Myrtaceae 


Olacaceae 


Passifloraceae 


Rosaceae 


Rutaceae 


Mangifera indica L. 


var. criollo 

var. manila 

var. Kent 
Spondias mombin L. 


Spondias purpurea L. 


Spondias radlkoferi Donn. Sm. 

Spondias sp. 

Tapirira mexicana Marchand 

Annona muricata L. 

Tabernaemontana alba Mill. 

Gonolobus niger (Cav.) R. Br. 

Quararibea funebris (La Llave) 
Vischer 

Carica papaya L. 

Terminalia catappa L. 

Ricinus communis L. 

Mammea americana L. 

Juglans pyriformis Liebm. 

Inga jinicuil G. Donht. 


Inga spuria Humbl. 

Inga vera Willd. 

Byrsonima crassifolia (L.) Kunth 

Trophis mexicana (Liebm.) Bur. 

Brosimum alicastrum Sw. 

Icacorea compressa (Kunth) Standley 

Myrciaria floribunda (H. West ex 
Willd.) O. Berg 

Psidium guajava L. 


Psidium guineense Sw. 


Psidium sartorianum (O. Berg.) Nied. 


Syzygium jambos (L.) Alston 
Ximenia americana L. 

Bunchosia biocellata Schlecht. 
Passiflora ciliata Aiton 

Passiflora edulis Sims. 

Passiflora edulis f. flavicarpa Deg. 
Cordia dodecandria Sesse & Moc. 
Crataegus mexicana DC. 
Eriobotrya japonica (Thunb.) Lindl. 
Prunus persica (L.) Batsch 
Casimiroa edulis La Llave & Lex. 
Citrus maxima (Burm.) Merr. 
Citrus reticulata Blanco 

Citrus paradisi Macfad. 

Citrus sinensis (L.) Osbek 


var. valencia 
var. navel 


Mango corriente 
Mango manila 
Mango petacon 
Jobo 


Ciruelo tropical 


Jobo Cimarron 
Cundoria 
Cacao silvestre 
Guanabana 
Huevo de gato 
Vaquitas 
Canela 


Papaya 
Almendro 
Higuerilla 
Zapote domingo 
Nogal 

Jinicuil 


Chalahuite peludo 
Chalahuite 
Nanche 

Jobo Cimarron 
Ramon (Ojite) 
Capulin silvestre 
Guayabilla 


Guayaba 


Guayaba acida 
Guayaba Tejon 
Pomarrosa 
Ciruela de monte 
Granada de arbol 
Granada roja 
Granada amarilla 
Maracuya 

Copite 

Tejocote 

Nispero 

Durazno 

Zapote Blanco 
Pomelo 
Mandarina 
Toronja 


Naranja Valencia 


Naranja Ombligona 


1-9, 11-12, 14-15, 17, 
19-21, 24, 27 


1-3, 5—6, 10, 16-17, 
19-21, 23, 24, 27-28 

1-6, 9-11, 13-17, 19— 
223 

24, 27 

3235.25 

27, 30 

5, 16-17 

10 

PONS PST 

10 


ts WS), 17 

Dy Sy MS 

21 

2 

34, 36 

2, 233 24, 20) 2 ee 
30, 34 

21, 24, 27 

21, 24, 27 


=H Nnnnu 

SEE Sse 
— 

by 

S) 


1-6, 9-17, 19-34, 36— 


21, 24, 27, 32, 36 
215 24.27 So 


2. 18: 235255355 
5; 6; LO sss OS iee2 ete 
26, 27, 29, 32, 34, 36 


VOLUME 102, NUMBER 4 


Table 2. Continued. 


Plant family Scientific name 


807 


Local common name Locality code* 


Citrus aurantium L. 


1, 9, 14-16, 19-21, 23- 
DD), AM, Xd SO), Dil. 


Naranja Cucha 


33, 40 
Sapotaceae Bumelia spiniflora A. DC. Pionche 17 
Calocarpum mammosum (L.) Zapote Mamey D2 23524 202i 
P. Royen 
Chrysophyllum mexicanum Zapote Nino 10 
Brandegee ex. Stand. 
Chrysophyllum cainito L. Caimito 8 
Manilkara zapota (L.) P. Royen Chico Zapote 14-17, 19 
Pouteria hypoglauca (Standl.) Baehni Zapote calentura DT 
Pouteria campechiana (Kunth) Baehni Zapote Nifio DY 
Solanaceae Solanum ionidium Bitter Tomatillo 36 
Staphyleaceae Turpinia insignis (H.B. & K.) Tul. Turpinia 36 


* Places in which plant species were sampled. Numbers correspond to those presented in Table 1. 


tured in McPhail traps in all four trapping 
sites. Toxotrypana curvicauda was captured 
only in Apazapan, Veracruz. 


DISCUSSION 


We discovered new host plants for An- 
astrepha bahiensis, A. hamata (2 plant spe- 
cies), A. chiclayae Greene, A. fraterculus, 
and most significantly, for the economical- 
ly-important species, A. obliqua. New host 
plants are also reported for Toxotrypana 
curvicauda, Rhagoletis zoqui, R. sp. and 
Hexachaeta sp. We were further able to 
document, for the first time, the presence of 
A. canalis, A. dentata, and A. limae Stone 
in the State of Veracruz, Mexico. 

Of the 17 species of Anastrepha reported 
by Hernandez-Ortiz (1992) in the state of 
Veracruz, we found all in fruits, or captured 
in McPhail traps. In addition, we also found 
the previously unrecorded A. alveata (Pie- 
dra et al. 1993), A. bicolor, A. canalis, A. 
dentata, and A. limae. 

Of the new records found for the genus 
Anastrepha, A. hamata stands out, not only 
because this is the first report of its host 
plants, but also because the larvae were 
found feeding only on seeds of two fruit 
species. Thus, this species is added to the 
list previously published by Hernandez-Or- 
tiz and Aluja (1993), in which A. cordata, 


A. sagittata (Stone), A. pallens Coquillett, 
A. crebra Stone, A. anomala Stone, and A. 
pickeli Lima are reported as seed-feeding 
species. We note however, that A. cordata 
and A. crebra also apparently feed in the 
mesocarp of fruits (Norrbom et al. 2000). 
Anastrepha bahiensis, A. fraterculus, and 
A. obliqua were found simultaneously at- 
tacking the previously unreported host Myr- 
ciaria floribunda (local common name: 
Guayabilla) (Myrtaceae), in the Apazapan 
region. We wonder if these species are un- 
able to recognize the host marking phero- 
mone of their congeners or if the extremely 
scarce supply of alternative host plants dur- 
ing the time of year when this plant is in 
fruit, causes flies to oviposit in previously 
utilized and marked fruit (due to a large eg- 
gload). For A. fraterculus and A. obliqua, it 
is relevant to mention that this plant can 
function as an alternative host from March 
until May. This could allow A. fraterculus 
to bridge the period when its previously re- 
ported hosts are unavailable (Psidium spp. 
prior to May, and S. jambos [L.] Alston 
from May to July). In the case of Anastre- 
pha obliqua, we believe that this host could 
also serve as a “‘bridge’’ between the fruit- 
ing periods of two wild hosts (Tapirira 
mexicana Marchand, which locally bears 
fruits from September to November, and 


808 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 3. Native and exotic host plants that harbored fruit flies in Veracruz, Mexico, during our nine year 
study (1990-1998). 


Common local name 


Scientific name 


Fruit fly species of host plant of host plant Plant family 
Anastrepha 
alveata Ciruela de monte Ximenia americana Olacaceae 
aphelocentema Zapote calentura Pouteria hypoglauca Sapotaceae 
bahiensis Guayabilla* Myrciaria floribunda Myrtaceae 
Ramon or Ojite Brosimum alicastrum Moraceae 
chiclayae Granada amarilla Passiflora edulis Passifloraceae 
Granada roja Passiflora ciliata Passifloraceae 
cordata Huevo de gato Tabernaemontana alba Apocynaceae 
crebra Canela Quararibea funebris Bombacaceae 
distincta Jinicuil Inga jinicuil Leguminosae 
Chalahuite peludo Inga spuria Leguminosae 
Chalahuite Inga vera Leguminosae 
fraterculus Almendro** Terminalia catappa Combretaceae 
Guayabilla Myrciaria floribunda Myrtaceae 
Guayaba Psidium guajava Myrtaceae 
Guayaba acida Psidium guineense Myrtaceae 
Guayaba tejon Psidium sartorianum Myrtaceae 
Pomarrosa Syzygium jambos Myrtaceae 
Durazno Prunus persica Rosaceae 
hamata Zapote Nino* Chrysophyllum mexicanum Sapotaceae 
Zapote Nino* Pouteria campechiana Sapotaceae 
ludens Mango Mangifera indica Anacardiaceae 
cultivar Criollo 
cultivar Manila 
cultivar Kent 
Durazno Prunus persica Rosaceae 
Zapote Blanco Casimiroa edulis Rutaceae 
Naranja cucha Citrus aurantium Rutaceae 
Pomelo Citrus maxima Rutaceae 
Toronja Citrus paradisi Rutaceae 
Mandarina Citrus reticulata Rutaceae 
Naranja dulce Citrus sinensis Rutaceae 
obliqua Mango Mangifera indica Anacardiaceae 
var. Manila 
var. Kent 
var. Criollo 
Ciruelo Spondias purpurea Anacardiaceae 
Cundoria Spondias sp. Anacardiaceae 
Jobo cimarron Spondias radlkoferi Anacardiaceae 
Jobo Spondias mombin Anacardiaceae 
Cacao Tapirira mexicana Anacardiaceae 
Guayabilla* Myrciaria floribunda Myrtaceae 
serpentina Zapote mamey Calocarpum mammosum Sapotaceae 
Zapote nino Chrysophyllum mexicanum Sapotaceae 
Caimito Chrysophyllum cainito Sapotaceae 
Chico Zapote Manilkara zapota Sapotaceae 
Zapote calentura Pouteria hypoglauca Sapotaceae 
striata Guayaba Psidium guajava Myrtaceae 
Guayaba acida Psidium guineense Myrtaceae 


Guayaba tejon 


Psidium sartorianum 


Myrtaceae 


VOLUME 102, NUMBER 4 


Table 3. Continued. 


809 


SsSesas"\VTQVWVWKK.® 5 


Common local name 


Fruit fly species of host plant 


Rhagoletis 
pomonella Tejocote 
turpiniae Turpinia 
sp. Tomatillo 
zoqui Nogal 


Hexachaeta 


(undescribed species) Jobo Cimarr6n* 


Toxotrypana 
curvicauda Papaya 


Vaquitas* 


Scientific name 


of host plant Plant family 


Rosaceae 
Staphyleaceae 
Solanaceae 
Junglandaceae 


Crataegus mexicana 
Turpinia insignis 
Solanum iodinium 
Juglans pyriformis 


Moraceae 


Trophis mexicana 


Caricaceae 
Asclepiadaceae 


Carica papaya 
Gonolobus niger 


* First report for Mexico. 
** First report for Veracruz State. 


Spondias purpurea L., available from April 
to June). Interestingly, M. floribunda is 
heavily preyed upon local mammals who 
eat practically every available fruit (trees 
bear fruit during the peak of the dry season 
and thus represent one of the few available 
food items for wildlife). Because of this, it 
is very difficult to find fruit in the field. 

The presence of A. fraterculus in fruits 
of Terminalia catappa corroborates the pre- 
vious report by Patino (1989) working in 
Papantla and Gutiérrez Zamora, Veracruz. 
However, we find it noteworthy that it was 
never found infesting any of the several Cit- 
rus species we sampled. 

Unfortunately, we were not able to rear 
A. bicolor, A. canalis, A. dentata, A. limae, 
A. pallens, A. robusta, A. spatulata, and A. 
zuelaniae from host fruits. These species 
were only captured in McPhail traps. How- 
ever, Hernandez-Ortiz (1992) pointed out 
that species belonging to the dentata group 
(e.g., A. dentata), daciformis group (e.g., A. 
bicolor and A. pallens) and robusta group 
(e.g., A. robusta) probably attack plants be- 
longing to the family Sapotaceae. Future ef- 
forts at discovering their host plants should 
thus be directed to all wild species of this 
family. According to Stone (1939) and Bak- 
er et al. (1944), A. pallens attacks Bumelia 
spiniflora A. DC. (Sapotaceae), while No- 
trbom (1998) pointed out that it infests 


fruits of Sideroxylon celastrinum (Kunth) 
T.D. Pennington and S. lanuginosa Michx. 
(Sapotaceae). We note that Norrbom (1998) 
considers that B. spiniflora was likely mis- 
spelled (the correct name should be B. spi- 
nosa A.DC.). Should the latter be true, then 
B. spiniflora is a synonym of S. celastrinum 
(Norrbom 1998). Anastrepha leptozona 
Hendel was also not reared from any of the 
fruits collected. However, Aluja et al. 
(1987) found it in Micropholis mexicana 
Gilly ex Cronquist (Sapotaceae), in the state 
of Chiapas, and Norrbom and Kim (1988) 
report its occurrence in 6 plant families. 
Norrbom et al. (1999) also note that A. zue- 
laniae, A. limae and A. canalis are associ- 
ated with plants in the families Flacourti- 
aceae, Passifloraceae and Staphylaceae, re- 
spectively. 

In the Los Tuxtlas region, Hernandez-Or- 
tiz and Pérez Alonso (1993) reported the 
presence of 13 species of Anastrepha. Of 
these, they hypothesized that A. crebra, A. 
minuta and Anastrepha sp. (close to A. per- 
dita) are apparently restricted in Mexico, to 
tropical, evergreen rain forests. Our results, 
indicating that these species are only pre- 
sent in the tropical decidous forests of cen- 
tral and northern Veracruz, lend support to 
this hypothesis. 

For Toxotrypana curvicauda, our report 
corroborates the previous records of Mason 


810 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 4. Fruiting phenology of fruit fly host plants in Veracruz, Mexico. 


Month 
Plant species J F M A M J J A S O N D 


Anastrepha host plants 


Brosimum alicastrum 4 
Calocarpum mammosum 


xx xX 
xx x 
x xX 
x 
* 


Casimiroa edulis 
Chrysophyllum mexicanum x xX 
Chrysophyllum cainito xX x 

Citrus aurantium 


* 
~ 


~ x 


Citrus maxima 
Citrus paradisi 


x MK xX 


Citrus reticulata 


xx KK XM 
* 
xx KK XK 
xa KKM 


Citrus sinensis 
Inga jinicuil Xx Xx »¢ 
Inga spuria 
Inga vera 
Mangifera indica 
cultivar Criollo 
cultivar Manila x 
cultivar Kent 
Manilkara zapota 
Myrciaria floribunda 
Passiflora edulis 
Passiflora ciliata 
Pouteria hypoglauca x x x 
Pouteria campechiana x x x 
Prunus persica x x xX xX 
Psidium guajava 
0-500* x Xx x x 
500—1,000* 
1 ,000—1,500* 
Psidium guineense 


~*~ xX 

xx KK 
xx K KK 
x x x x 
x x* 
x mK KM 

x mK x 

~*~ x 


~*~ 
* 


x x 
~ x 
xxx 


Psidium sartorianum 
Quararibea funebris 
Spondias purpurea x x x x 

Spondias mombin xX x 
Spondias radlkoferi x 
Spondias sp. Xx x 
Syzygium jambos x 
Tabernaemontana alba 


xx x 
x x*K xX 
xx mK mK MK 
x mex MK 
xx 


x x 

~*~ x 
nK % 
* 


Tapirira mexicana 
Terminalia catappa x xX xX x 
Ximenia americana x x x 


x xK x 
~*~ x 


Hexachaeta host plants 
Trophis mexicana x xX 

Toxotrypana host plants 
Carica papaya xX x x x x xX xX xX 
Gonolobus niger x xX x 

Rhagoletis host plants 


Crataegus mexicana 


x x 
~*~ 


Juglans pyriformis 
Solanum ionidium 


mM XM 
ax KK 


Turpinia insignis 


* Meters above sea level. 


VOLUME 102, NUMBER 4 


S316) 


300 


2S0 


200 


100 


MEAN NUMBER OF LARVAE/KG. OF FRUIT 
ul 
Oo 


@) 100 


300 


811 


400 500 700 


FRUIT WEIGHT 


Fig. 1. 


(1922), Castrej6n-Ayala and Camino-Lavin 
(1991), and Landolt (1994), who pointed 
out that adults of this species feed mainly 
on fruits belonging to the families Carica- 
ceae and Asclepiadaceae. Here we report 
this species infesting Gonolobus niger, a 
plant also belonging to the family Ascle- 
piadaceae. 

For Hexachaeta sp., we are the first to 
report its host plant as Trophis mexicana. 
This confirms that various species belong- 
ing to this fly genus develop in fruits of the 
Moraceae. 

We are able to report a new host plant 
(Juglans pyriformis) for R. zoqui. The only 
other known host plant record for this fly 
species is J. mollis Engelm. (Bush 1966). 
Further, we also report for the first time that 
Solanum ionidium is a host plant of an un- 
described species in the genus Rhagoletis. 
This adds further evidence about the im- 
portance of the family Solanaceae as hosts 
of this genus. 

The polyphagy observed in A. ludens (8 
plant species belonging to 3 families), A. 


Number of larvae per kilogram of fruit as influenced by size of fruit. 


obliqua (7 plant species belonging to 2 
families) and A. fraterculus (7 species be- 
longing to 4 families) has previously been 
discussed by Aluja et al. (1987) and Her- 
nandez-Ortiz and Aluja (1993). In our opin- 
ion, this polyphagy permits these four spe- 
cies to be abundant during practically the 
whole year. Anastrepha bahiensis was also 
found to be a polyphagous species, with lar- 
vae feeding on fruits belonging to two gen- 
era in two different families (M. floribunda 
[Myrtaceae] and Brosimum alicastrum Sw. 
[Moraceae]). 

In contrast, some species in our study ap- 
peared to be monophagous. This is the case 
for A. alveata feeding on Ximenia ameri- 
cana (Olacaceae) (Piedra et al. 1993), and 
A. aphelocentema on Pouteria hypoglauca 
(Sapotaceae). Anastrepha aphelocentema 
was found only in the northern part of Ve- 
racruz (Papantla) (Patifio, 1989), in spite of 
the fact that this plant is also present in lo- 
calites such as La Mancha, approximately 
150 km away from Papantla. In La Mancha, 
fruits were attacked only by A. serpentina. 


812 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 5. Infestation patterns by Anastrepha flies based on size of host fruit. 


eT 


Degree of 
infestation 
Total no. Total no. Fruit fly (larvae/kg 
Host plant Kg. sampled of pupae of adults species of fruit) 
Large (300—1,000 gr) 
Calocarpum mammosum 143.73 3,942 1, TSI A. serpentina 27.43 
Citrus paradisi 1,345.71 22,739 19,428 A. ludens 16.90 
Mangifera indica 
cultivar Kent 56.57 902 ES95 
27 A. obliqua 
716 A. ludens 
Citrus maxima 79.66 853 S52 A. ludens 10.70 
Carica papaya 126.50 529 477 T. curvicauda 4.18 
Medium (50-300 gr) 
Chrysophyllum cainito 11.00 2,210 1,926 A. serpentina 201.00 
Prunus persica 10.62 338 31.83 
228 A. fraterculus 
Si) A. ludens 
Manilkara zapota 101.83 3,178 Dab A. serpentina 31.21 
Pouteria hypoglauca? 19.05 506 328 A. aphelocentema 26.56 
Casimiroa edulis B95) 952 477 A. ludens 24.10 
Pouteria hypoglauca' 17.28 369 Dray A. serpentina 21.35 
Juglans pyriformis WL ES aV7/ 899 R. zoqui 20.51 
Citrus sinensis 264.27 Spyd57/ 2,742 A. ludens 20.27 
Citrus aurantium 669.29 1S F503 525 A. ludens 20.20 
Mangifera indica 
var. criollo 121.78 eZ 14.55 
20 A. ludens 
995 A. obliqua 
Citrus reticulata 2.80 36 25 A. ludens 12.86 
Chrysophyllum mexicanum 59.36 531 342 A. hamata 8.94 
Mangifera indica 
var. manila 42.97 337 7.84 
58 A. ludens 
222 A. obliqua 
Terminalia catappa 10.09 55 10 A. fraterculus 5.45 
Inga jinicuil 22.38 Vi 64 A. distincta 3.44 
Small (<5O gr) 
Spondias mombin 64.67 28,138 4,241 A. obliqua 435.10 
Ximenia americana 16.08 4,960 2,082 A. alveata 308.46 
Turpinia insignis 12225 3,416 674 R. turpiniae 278.90 
Psidium guajava 97.60 16,619 170.28 
4,879 A. striata 
8,923 A. fraterculus 
Spondias sp. 29.20 4,514 2ASS A. obliqua 154.59 
Inga vera 0.56 76 76 A. distincta PS S2k 
Psidium guineense 4.76 509 106.93 
ie A. striata 
359 A. fraterculus 
Solanum ionidium 0.29 29 11 R. sp. 100.00 
Psidium sartorianum 3.62 282 77.90 
47 A. striata 
43 A. fraterculus 
Tapirira mexicana 14.93 1,060 325 A. obliqua 71.00 
Spondias radlkoferi TST 384 131 A. obliqua 50.72 


nn 


VOLUME 102, NUMBER 4 813 

Table 5. Continued 

Degree of 

infestation 

Total no. Total no. Fruit fly (larvae/kg 

Host plant Kg. sampled of pupae of adults species of fruit) 

Inga spuria 10.52 495 373 A. distincta 47.05 

Crataegus mexicana 22.50 986 559 R. pomonella 43.83 

Spondias purpurea 182.34 6,170 ils} A. obliqua 33.84 

Passiflora ciliata 0.84 24 18 A. chiclayae 28.57 

Syzygium jambos DN (9) 615 389 A. fraterculus 28.01 

Quararibea funebris 3.26 89 67 A. crebra 2-30 

Trophis mexicana 0.21 5) 72 Hexachaeta sp. 23.81 

Passiflora edulis 0.67 10 A. chiclayae 14.93 

Tabernaemontana alba 16.43 62 49 A. cordata Seyi 

Gonolobus niger 5.00 “3 ce T. curvicauda — 

Brosimum alicastrum ca. 0.30 “A A. bahiensis — 

Myrciaria floribunda ca. 0.30 2 A. obliqua — 

A. fraterculus 
A. bahiensis 


' Fruit collected in La Mancha. 
? Fruit collected in Morgadal (Poza Rica). 
* Pupae and adults that emerged were not counted. 


Table 6. Fruit fly species captured in McPhail traps 
in Central Veracruz, Mexico. 


Llano Monte M. de la 
Fruit fly species Apazapan Grande Blanco Torre 
A. alveata xX xX xX 
A. bahiensis x 
A. bicolor xX xX xX xX 
A. canalis xX 
A. chiclayae xX x x xX 
A. cordata xX 
A. dentata xX 
A. distincta xX x 4 xX 
A. fraterculus 4 Xx Xx XxX 
A. hamata xX xX 
A. leptozona x Xx 
A. limae xX »4 
A. ludens x xX xX xX 
A. obliqua x x x x 
A. pallens Xx 4 
A. robusta xX 
A. serpentina xX x x x 
A. spatulata x xX Xx X 
A. striata xX xX xX 
A. zuelaniae Xx 
T. curvicauda xX 
Hexachaeta spp. Xx Xx x Xx 


It is possible that the recent heavy habitat 
alteration caused local extinction of this 
species. 

It is significant that in the Monte Blanco 
region (central Veracruz) fruits of Pouteria 
campechiana were attacked only by A. ha- 
mata (on seeds), whereas in Los Tuxtlas 
(southern part of the state), Chrysophyllum 
mexicanum fruits were attacked by both A. 
serpentina on pulp, and A. hamata on 
seeds. It is noteworthy that two species of 
Anastrepha can utilize the same host with- 
out competition. Pouteria campechiana 
seeds and pulp were also reported attacked 
by both A. sagittata and A. serpentina, re- 
spectively, by Baker et al. (1944). However, 
Aluja et al. (1987) reported only A. serpen- 
tina larvae feeding on pulp of P. campe- 
chiana fruits in the State of Chiapas (Mex- 
ico), but no trace of A. sagittata. 

In other Anastrepha species, we found 
that their host plants belonged to multiple 
species of only one genus (stenophagous 
species). This was true for A. distincta in 
Inga spuria Humbl. and J. vera Willd. (Leg- 
uminoseae), for A. striata Schiner in Psi- 
dium guajava L., P. guineense Sw. and P. 
sartorianum (O. Berg) Nied. (Myrtaceae) 


814 


and for A. chiclayae in Passiflora edulis 
Sims. and P. ciliata Aiton (Passifloraceae). 
We also confirm that A. serpentina is an 
oligophagous species with larvae found 
feeding in four fruit species belonging to 
the Sapotaceae (see Hernandez-Ortiz 1992). 
Interestingly, A. hamata was also found to 
be an oligophagous species (larvae feed on 
seeds of 2 plant species belonging to the 
Sapotaceae). 

Even though trapping efforts were made 
in an extended region of central Veracruz, 
A. dentata, A. robusta, and A. zuelaniae 
were captured only in Apazapan, where the 
vegetation has been characterized as low 
deciduous forest. In contrast, adults of A. 
canalis were captured only in Monte Blan- 
co, where the vegetation was originally 
comprised of montane cloud forest. Anas- 
trepha limae was found in both Llano 
Grande and Monte Blanco. This suggests 
that the geographic distribution of some An- 
astrepha species is restricted to certain 
zones, possibly because their host plants are 
present only in these zones. 

We also found that the timing of capture 
of certain monophagous species coincided 
with the fruiting periods of their host plants. 
For example, A. hamata and A. alveata 
adults were captured in McPhail traps al- 
most exclusively when their host fruits 
were available (P. campechiana and C. 
mexicanum, and X. americana, respective- 
ly). In our opinion, this is probably due to 
an extraordinary capacity of adult survival 
for long periods of time. Under laboratory 
conditions, A. alveata adults can live up to 
431 days (Aluja and Jacome, unpublished 
data). Such adult longevities probably allow 
adults to lay eggs in the fruits produced the 
following year. However, the locations 
where the adults of such species persist dur- 
ing most of the year are unknown. 

Finally, our data on the relationship of 
fruit size and larval infestation levels rep- 
resents further evidence that small, wild 
fruits are more heavily infested than larger, 
commercially grown fruits that were for the 
most part, recently (on an evolutionary 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


scale) introduced to the Americas. This re- 
lationship was recently discussed by Aluja 
et al. (2000) who indicated that native flies 
have probably still not fully developed the 
ability to metabolize the toxic chemicals 
that these exotic plants contain. As a result, 
and despite the fact that females lay many 
eggs in them, few eggs hatch or most larvae 
die or develop poorly. 


ACKNOWLEDGMENTS 


We thank Jestis Reyes Flores for having 
encouraged us to do this study, Vicente 
Hernandez-Ortiz for identifying all fly spe- 
cies reported here, and Isabel Jacome, Anita 
Sanchez, Braulio Cérdoba, Guadalupe Tru- 
jillo, Gloria Lagunes, and Anabel Davila 
(all Instituto de Ecologia, A.C., Xalapa, Ve- 
racruz, Mexico) for technical assistance. We 
also thank Leticia Lagunes, Pedro Cérdoba, 
Augusto Martinez, Faustino Cabrera and 
the Bigurra-Armida family for allowing us 
to collect fruits and install McPhail traps in 
their orchards. Thanks are also due to Car- 
los Duran, and Sergio Avendano (Herbari- 
um IXAL, Instituto de Ecologia, A.C.) for 
identifying plant material and to Diana 
Pérez-Staples for reviewing an early ver- 
sion of the manuscript. Useful critical re- 
views were also made by A.L. Norrbom 
and an anonymous referee. Partial finan- 
cial support was provided by the Consejo 
Nacional de Ciencia y Tecnologia 
(CONACyT) (Grant D111-903537), the 
Comisi6n Nacional para el Conocimiento y 
Uso de la Biodiversidad (CONABIO) 
(Grant FB325/H296/96), the United States 
Department of Agriculture-Agricultural Re- 
search Service (Agreement 58-6615-3- 
025), the Sistema Regional de Investigaci6n 
del Golfo de México (SIGOLFO) (Grant 
96-01-003-V) and the Campafia Nacional 
Contra las Moscas de la Fruta Mexico Se- 
cretaria de Agricultura, Ganaderia y Desar- 
rollo Rural—Instituto Interamericano de 
Cooperacion para la Agricultura (SAGAR- 
IICA). 


VOLUME 102, NUMBER 4 


LITERATURE CITED 


Aluja, M. 1994. Bionomics and Management of An- 
astrepha. Annual Review of Entomology 39: 
151-174. 

Aluja, M., J. Pinero, I. Jacome, E Diaz-Fleischer, and 
J. Sivinski. 2000. Behavior of flies in the genus 
Anastrepha (Trypetinae: Toxotrypanini), pp. 375— 
406. In Aluja, M. and A.L. Norrbom, eds., Fruit 
Flies (Diptera: Tephritidae): Phylogeny and Evo- 
lution of Behavior. Boca Raton, FL, USA. CRC 
Press, 944 pp. 

Aluja, M., J. Guillén, G. de la Rosa, M. Cabrera, H. 
Celedonio-Hurtado, P. Liedo, and J. Hendrichs. 
1987. Natural host plant survey of the economi- 
cally important fruit flies (Diptera: Tephritidae) of 
Chiapas, Mexico. Florida Entomologist 70: 329— 
338. 

Aluja, M., H. Celedonio-Hurtado, P. Liedo, M. Ca- 
brera, E. Castillo, J. Guillén, and E. Rios. 1996. 
Seasonal population fluctuations and ecological 
implications for management of Anastrepha fruit 
flies (Diptera: Tephritidae) in commercial mango 
orchards in Southern Mexico. Journal of Econom- 
ic Entomology 89: 654—667. 

Baker, A. C., A. C. Stone, C. C. Plummer, and M. 
McPhail. 1944. A review of studies on the Mex- 
ican fruit fly and related Mexican species. USDA- 
APHIS Miscellaneous Publication No. 531. 154 
Pp. 

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 (Harvard University) 134: 
431-562. 

Castrejon-Ayala, F and M. Camino-Lavin. 1991. New 
host plant record for Toxotrypana curvicauda 
(Diptera: Tephritidae). Florida Entomologist 74: 
466. 

Hernandez-Ortiz, V. 1992. El género Anastrepha Schi- 
ner en México (Diptera: Tephritidae). Taxonomia, 
distribucion y sus plantas hospederas. Instituto de 
Ecologia Publ. No. 33, Xalapa, Veracruz, México, 
162 pp. 

Hernandez-Ortiz, V. and M. Aluja. 1993. Listado de 
especies del genero neotropical Anastrepha (Dip- 
tera: Tephritidae) con notas sobre su distribucion 
y plantas hospederas. Folia Entomologica Mexi- 
cana 88: 89-105. 

Hernandez-Ortiz, V. and R. Pérez-Alonso. 1993. The 


815 


natural host plants of Anastrepha (Diptera: Te- 
phritidae) in a tropical rain forest of Mexico. Flor- 
ida Entomologist 76: 447—460. 

Landolt, P. J. 1994. Mating frequency of the papaya 
fruit fly (Diptera: Tephritidae) with and without 
host fruit. Florida Entomologist 77: 305-312. 

Mason, A. C. 1922. Biology of the papaya fruit fly 
Toxotrypana curvicauda in Florida. U.S. Depart- 
ment of Agriculture Bulletin No. 1081, 10 pp. 

Norrbom, A. L. 1998. A revision of the Anastrepha 
daciformis species group (Diptera: Tephritidae). 
Proceedings of the Entomological Society of 
Washington 100: 160-192. 

. 2000. Host plant database for Anastrepha and 
Toxotrypana (Diptera: Tephritidae: Toxotrypani- 
ni), Diptera Data Dissemination Disk 2. 

Norrbom, A. L. and K. C. Kim. 1988. A list of the 
reported host plants of the species of Anastrepha 
(Diptera: Tephritidae). USDA-APHIS Miscella- 
neous Publication No. 81-52, 114 pp. 

Norrbom, A., L. E. Carroll, FE C. Thompson, I. M. 
White, and A. Freidberg. 1999. Systematic Data- 
base of Names, pp. 65-251. Jn Thompson, E C., 
ed., Fruit Fly Expert Identification System and 
Systematic Information Database. Myia 9, 524 pp. 

Norrbom, A., R. A. Zucchi and V. Hernandez-Ortiz. 
2000. Phylogeny of the Genera Anastrepha and 
Toxotrypana (Trypetinae: Toxotrypanini) Based 
on Morphology. pp. 343-362. In Aluja, M. and A. 
Norrbom, eds., Fruit Flies (Diptera: Tephritidae): 
Phylogeny and Evolution of Behavior. CRC Press. 
Boca Raton, Florida, 944 p. 

Patino, J. 1989. Determinacion de las especies de An- 
astrepha Schiner (Diptera: Tephritidae) en frutales 
y citricos de Papantla y Gutiérrez Zamora, Vera- 
cruz, Bsc. Thesis, Universidad Veracruzana, Tux- 
pan, Veracruz, México. 

Piedra, E., A. Zuniga, and M. Aluja. 1993. New host 
plant and parasitoid record in Mexico for Anas- 
trepha alveata Stone (Diptera: Tephritidae) Pro- 
ceedings of the Entomological Society of Wash- 
ington 95: 127. 

Stone, A. 1939. A new genus of Trypetidae near An- 
astrepha (Diptera). Journal of the Washington 
Academy of Science 29: 340—350. 

USDA, ARS. 2000. National Genetic Resources Pro- 
gram. Germoplasm Resources Information Net- 
work—(GRIN). [Online Database] National Germ- 
plasm Resources Laboratory, Beltsville, Mary- 
land. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 816-830 


REVISION OF THE GENUS PARADA HORVATH (HEMIPTERA: TINGIDAE) 
WITH CLADISTIC ANALYSIS 


ErIc GUILBERT 


ESA 8043 du CNRS, Museum National d’Histoire Naturelle, 


Laboratoire 


d’Entomologie, 45 rue Buffon, F-75005 Paris, France (e-mail: guilbert@mnhn.fr) 


Abstract.—Three new species of Parada from New Caledonia are described: P. minuta, 
P. paitae and P. majuscula. The monotypic Alloeocysta approba Drake is considered a 
junior synonym of Parada and to which its only included species is transferred. The 
evolution of certain characters, and the relationships among species of Parada are pre- 
sented together with a cladistic analysis. An identification key for these species is also 


given. 


Key Words: 


Horvath (1925) erected Parada as a sub- 
genus of Cysteochila Stal for the reception 
of Cysteochila (Parada) taeniophora Hor- 
vath (1925). Cysteochila has the paranota 
widely reflected, inflated, covering the lat- 
eral carinae, and the internal margins (« mar- 
gine interno ») rounded, while Parada has 
the paranota narrowly reflexed, superficially 
adpressed onto the pronotum and not cov- 
ering the lateral carinae, and the internal 
margins slightly sinuate and never rounded 
(Horvath 1925). Drake (1942) raised Par- 
ada to generic level, and add two new spe- 
cies to the genus (P. torta and P. popla). 
Later, he added four more new species to 
the genus (P. absona, P. darlingtoni and P. 
hackeri Drake 1952; P. solla Drake and Ru- 
hoff 1961). 

The genus presently comprises seven 
Australian species (New South Wales and 
Queensland) (Drake and Ruhoff 1965). 
Three new species from New Caledonia are 
described here. In addition, Drake (1961) 
described a new genus and a new species 
from a single specimen caught in New 
South Wales: Alloeocysta approba. This 
species, the only included species of the ge- 


Parada, Tingidae, new species, evolution, phylogeny, key 


nus, shares with Parada the form of the lat- 
eral carinae, the paranota and the hood, and 
is considered a junior synonym. Alloeocysta 
approba Drake is transferred to Parada. In 
addition, Cysteochila cubens Guilbert (in 
press), described from New Caledonia 
shows affinities with the species of Parada 
and could be an intermediate species be- 
tween these two genera. Among these re- 
lationships, the evolution of some charac- 
ters getting narrower are discussed through 
a cladistic analysis. Deposition of types is 
specified: MNHN for Muséum National 
d’Histoire Naturelle, Paris; BPBM for 
Bernice P. Bishop Museum, Hawaii. All 
measurements are in millimeters. 


SYSTEMATICS 
Parada Horvath 1925 


Cysteochila (Parada) Horvath 1925: 3. 
Type species: Cysteochila (Parada) tae- 
niophora Horvath 1925: 3, by orig. desig. 

Parada: Drake 1942: 4. 

Alloeocysta Drake 1961: 109. New Syn- 
onymy. Type species: Alloeocysta appro- 
ba Drake 1961: 109, by orig. desig. 


VOLUME 102, NUMBER 4 


Diagnosis.—Head with at least two fron- 
tal and two occipital spines. Bucculae wide, 
with apices meeting in front. Antenna long, 
slender, first segment short, second small- 
est, third longest and fourth longer than first 
two together, pubescent. 

Pronotum gibbose, deeply punctuate, are- 
olate at apex, pubescent near collar, tricar- 
inate. Median carina long, slightly erected, 
contiguous to hood. Lateral carinae elevat- 
ed, ending before collar, curved inward, not 
resting on pronotum, modified as to form a 
long inflated areolate cyst. Hood inflated, 
raised. Paranota long, wide, reflexed up- 
right, not resting on pronotum, three to four 
areolae broad, almost reaching lateral cari- 
nae. 

Hemelytra wider and longer than abdo- 
men, slightly wider than pronotum width. 
Costal area uni- to biseriate, outer margins 
bent upward. Subcostal area bi- to triseriate, 
outer margins bent downward. Discoidal 
area five to seven areolae broad. Sutural 
area large, with larger areolae than subcos- 
tal and discoidal areas. Hypocostal laminae 
uniseriate. 


Parada approba (Drake), new 
combination 
(Figs. 7—8) 


Alloeocysta approba Drake 1961: 109. 


Comments.—According to Drake (1961) 
Alloeocysta approba is different “from oth- 
er Australian genera by having the hood 
distinctly wider than long and the lateral ca- 
rinae inflated, cyst-like on disc of prono- 
tum’”’ (Holotype measurements, body length 
2.99; body width 1.26; hood length 0.37; 
hood width 0.55). All species of Parada 
have lateral carinae inflated like A. approba 
and some of them have a hood wider than 
long. This is the case of P. majuscula and 
P. paitae. Alloeocysta approba has anten- 
nae as long as that of the species of Parada 
mesOA5: I;.0:12: Ill, 0.85: IV, 0.3); a pron- 
otum gibbose, deeply punctuate and areo- 
late at apex, a hood forming a vesicule wid- 
er than long, lateral carinae modified as to 


817 


form a subcylindrical areolate cyst that is 
closed inward by the internal margins 
touching the pronotum, while it is not the 
case for the species of Parada. The hem- 
elytra are a little longer than abdomen with 
a costal area uniseriate, a subcostal area bis- 
eriate and a discoidal area four to five are- 
olae broad. Despite some characters which 
distinguish A. approba from species of Par- 
ada; approba being the unique species of 
the genus Alloeocysta, the latter is consid- 
ered a synonym of the genus Parada. Thus, 
I propose the new combination Parada ap- 
proba (Drake) n. comb. 

Type.—Holotype: ¢, Bogan river, New 
South Wales, Australia (Australian Muse- 
um, Sydney). 


Parada majuscula Guilbert, new species 
(Figs. 1-3) 


Description.—Head and body black, legs 
and antenna yellowish to brown fuscous, 
distal part of fourth antennal segment and 
tarsi brown to black. Pronotum clear brown 
to fuscous. Hemelytra yellowish with three 
dark brown spots, one near base that may 
occupy just basal third of costal area or be 
expanded to cover all of the basal third of 
the hemelytron, plus clear brown spots at 
apical third and apex of costal area. Length 
3.05; width 1.17. 

Head with two frontal and two small, 
slender occipital spines. Bucculae wide, 
triseriate, with apices meeting in front. La- 
bium reaching hind coxae and labial sulcus 
sinuate. Sternal laminae widely separated 
on metasternum. Antennal measurements: I, 
0.192 I, 0.12: If, 1.01: IV, 045).. last an- 
tennal segment pubescent. 

Pronotum gibbose, deeply punctuate, are- 
olate on hind process, pubescent near collar, 
tricarinate. Median carina long, slightly el- 
evated, contiguous to hood. Lateral carinae 
elevated, ending before collar, curved in- 
ward, not resting on pronotum, modified as 
to form a long inflated open cyst, four are- 
olae broad. Hood inflated, raised, slightly 
wider than long (length 0.33; width 0.38). 
Paranota long, wide, reflexed and partially 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


818 


1 mm. 


Parada majuscula, habitus. Scale 


orale 


VOLUME 102, NUMBER 4 


TON 
ac CaS ee 


819 


? SS 


Figs. 2-3. 


recurved above pronotum, not resting on 
pronotum, four small areolae broad, almost 
reaching lateral carinae. 

Hemelytra wider, longer than abdomen, 
slightly wider than pronotum width. Costal 
area bent upward, mostly biseriate but un- 
iseriate at apex, outer areolae larger than 
inner. Subcostal area bent downward, tris- 
eriate. Discoidal area pentaseriate. Sutural 
area large, with larger areolae than subcos- 
tal and discoidal areas at apex. Hypocostal 
laminae uniseriate. 

Types.—Holotype: 36, New Caledonia, 
Riviere Bleue P7, dense evergreen rainfo- 
rest, 24.X.1992, L. Bonnet de Larbogne, J. 
Chazeau & E. Guilbert colls (fogging) 
(MNHN). Paratypes: 1 2, same data as ho- 
lotype (MNHN); 3 6 and 1 2, New Cale- 
donia, Yahoue III.1978, N.L.H. Krauss 
coll., Acc. #1978.114 (BPBM); 1 2°, New 
Caledonia, 6 km N of Paita, 25.1.1963, 


Parada majuscula. 2, Profile. 3, Ventral face. Scale = 1 mm. 


C.M. Yoshimoto coll. (BPBM); 1 ¢, New 
Caledonia, Mt Koghis, 400—600 m, II.1980, 
N.L.H. Krauss coll., Acc. #1980.128 
(BPBM). 

Comments.—The body length of this 
species may be sexually dimorphic (Table 
1), but this characteristic cannot be assessed 
here because of the small number of spec- 
imens. 


Parada paitae Guilbert, new species 
(Figs. 4—5) 


Description.—Head and body brown. 
Pronotum, hemelytra, legs and antenna 
clear to yellowish, apex of last antennal 
segment brown fuscous. Length 2.98; width 
1.17; hood length 0.31; hood width 0.31. 

Same characters as P. majuscula, except 
rostrum extends between median coxae but 
does not reach hind coxae, costal area is 
uniseriate, areolae are rather large, except 


820 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 1. Means of body, hood and antennal segments measurements for 5 males and 3 females of P. ma- 
Juscula in mm. 


el 
Body Hood 


Antennal Segments 
Length Width 


Length Width I II Il IV 
em 
Males POON 1.102 0.32 0.372 1.003 0.366 0.191 0.123 
Females 3.149 L27Z 0.349 0.400 1.010 0.292 0.200 0.123 
Both 3205 1.165 0.331 0.383 1.006 0.451 0.194 0.123 


O83 


wee 


SS 


SV / 
4 


S 
< 
ao 


Figs. 4-6. Parada spp. 4, P. paitae, right hemelytra, paranota and lateral carina of the habitus. 5, P. paitae, 
ventral face. 6, P. minuta, right hemelytra, paranota and lateral carina of the habitus. Scale = 1 mm. 


VOLUME 102, NUMBER 4 


821 


Body length, body width, hood length, hood width and antennal segments measurements of the 


New Caledonian species and P. approba. Last antennal segment measurements on P. approba is missing on the 


Table 2. 
type. 
Body 

Species Length Width Length 
Parada majuscula 3.05 oJ 0.33 
Parada paitae 2.98 7 0.31 
Parada minuta 2.63 0.94 0.29 
Parada approba 2.99 1.26 0.37 


one or two areolae divided into two areolae 
at level of apex, and discoidal area five to 
six areolae broad. Antennal segments mea- 
surements: I, 0.17: II, 0.12: III, 0.94: IV, 
0.40. 

Types.—Holotype: 2°, New Caledonia, 
Mont Nondoué near Paita, sclerophyllous 
forest, 16.X.1992, L. Bonnet de Larbogne, 
J. Chazeau & E. Guilbert colls (fogging) 
(MNHN). Paratype: 1 2, New caledonia, 
Mont Nondoué near Paita, sclerophyllous 
forest, 28.1V.1993, L. Bonnet de Larbogne, 
J. Chazeau & E. Guilbert colls (fogging) 
(MNHN). 


Parada minuta Guilbert, new species 
(Fig. 6) 


Description.—Head and _ pronotum 
brown, body, hemelytra, legs and antenna 
clear to yellowish, last antennal segment 
brown fuscous, some brown spots on hem- 
elytra on discoidal and sutural areas. Length 
2.63; width 0.94; hood length 0.29; hood 
width 0.31. 

Head short, with four tiny spines. Anten- 
nal segments as two former species. Anten- 
nal segments measurements: I, 0.15: II, 
0.11: Il, 0.92: IV, 0.43. Bucculae short, 
wide, three areolae broad. Labium short, 
not extending beyond mesosternum, labial 
sulcus widened between hind coxae. 

Pronotum gibbose, areolate, tricarinate, 
median carina uniseriate, elevated, lateral 
carinae wide, four areolae broad. Paranota 
four areolae broad. 

Costal area completely uniseriate, sub- 
costal area biseriate, discoidal area short, 


Hood 


Antennal Segments 
Width I II Ii IV 
0.38 0.19 OM 1.01 0.45 
0.31 0.17 0.12 0.94 0.40 
0.31 0.15 0.11 0.92 0.43 
0.55 0.15 0.12 0.85 — 


half length of hemelytra, five areolae broad 
at widest part. 

Types.—Holotype: 6, New caledonia, Ile 
des Pins: Vao, 0-100 m, I. 1985, N.L.H. 
Krauss coll. (BPBM). Paratype: 1 6, same 
data as the holotype (BPBM). 


KEY TO SPECIES OF PARADA 


1. Head armed with 4 cephalic spines....... D 
Head armed with 5 cephalic spines....... 4 
2. Subcostal area triseriate at widest part .... 3 


— Subcostal area biseriate at widest part (Fig. 6) 

P. minuta Guilbert, n. sp. 

3. Labium extending beyond hind coxae (Fig. 3) 
Berend eo acee nines okra P. majuscula Guilbert, n. sp. 

— Labium not reaching hind coxae (Fig. 5)... 

P. paitae Guilbert, n. sp. 

4 Paranotasthreesateolac wide ys acne es 5 


Paranota four areolae wide 
5. Median carina areolate and uniseriate (Fig. 
IES) Loeieaeeai erica ieR Ceol P. solla Drake and Ruhoff 
=) Medianvcarina not arcolates; - ere 
el eevee eee he kas P. taeniophora (Horvath) 
6. Lateral carinae five areolae wide (Fig. 10) . . 
MERE SARS Cons RR Oto ore P. absona Drake 
Lateral carinae less than five areolae wide .. 7 
7. Third antennal segment 4 times longer than 
last = .. yelePe MAE eee dae:s P. approba (Drake) 
— Third antennal segment less than 4 times lon- 
gerthan lasts. aa aS ioe w cee Bets 8 
8. Costal area with areolae the same size than 
that of subcostalearcay(sea 2) eee 
Pace ina ema ld Spee a te iE P. hackeri Drake 
— Costal area with areolae larger than that of 
subcostalearearanan = cries on eee 9 
9. Lateral carinae width of 2 areolae broad (Fig. 
Aa Aa deas de one eck P. darlingtoni Drake 
— Lateral carinae width of 3 areolae broad .. 10 
10. Costal area largely biseriate at base (Fig. 
P. popla Drake 
— Costal area uniseriate except for two areolae 


across) very base (Fis: I) S44 3 P. torta Drake 


OWY 
Gots) SLO « 


S8seoosg © A iS Ss 


Figs. 7-8. 


CLADISTIC ANALYSIS 


Choice of the taxa.—The eleven species 
of the genus Parada are included in this 
analysis as the ingroup. Three species are 
included as outgroups: Cysteochila cubens 
Guilbert, Physatocheila dissimilis Guilbert 
and Nobarnus pilosus Guilbert. Cysteochila 
cubens shows some affinities with the genus 
Parada, while N. pilosus is clearly different. 
Physatocheila dissimilis could be considered 
as intermediate between these two genera by 
its general morphology. Therefore, 14 spe- 
cies were included in this analysis. 

Character states.—Sixty-one morpholog- 
ical characters are analysed. They are ana- 
lysed in two differents ways. A first anal- 
ysis concern all the characters. A second 
analysis concern only fifty-one characters 


% ootssoeO Ose 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Parada approba. 7, Profile. 8, Ventral face. Scale = 1 mm. 


active and ten inactive. These ten characters 
define the size of various structures in terms 
of number of areolae. They are not used in 
the analysis because the homology between 
the areolae cannot be established with cer- 
tainty, but they figure as attributes. When a 
structure has two areolae, does it mean that 
an areola divided into two areolae, or does 
a second areola appears near the existing 
one? The results of the two analyses are 
compared. Inactive characters in the second 
analysis are mentioned. Characters of the 
genitalia are not taken into account because 
they do not vary within the genus. Multi- 
state characters are treated unordered. The 
matrix of characters is given in Table 3. 
Question marks represent characters lacking 
or uncertainly stated. 


VOLUME 102, NUMBER 4 823 


; Sa 
>) 
We’ 
SS 
iy 
we 
3 


4 DOD AS) 
fi Noo re is 
MS) wee °0) Ves 
; 4 A 7 rf} i 
dj 0 \\ | 
, ; go \ pg 
Us 6 U ladl \ 
(y S D i 
0); g @ 1h 
i S toy \ | 
) | \0 
D 0) © \ Q 
0 ic) \9 ) 0 
© \ 0 
\ 9 Ka Q 0 
i) a ic ; a 
0 5 \ ; ¢ 
Q 6) 
by 4 " D 
f ; Y \ : 
a () Q Xo =: 9) 
“0 C Se \ 
Q ° Xe 
9 \S 10 11 12 Xe 


Figs. 9-15. Parada spp. left costal area, paranota and lateral carinae. 9, P. popla. 10, P. absona. 11, P torta. 
12, P. hackeri. 13, P. taeniophora. 14, P. darlingtoni. 15, P. solla. Scale = 1 mm. 


0. Head black (0), brown (1), yellowish to 12. Occipital spines stout (0), slender (1). 


white (2). 13. Median spine long (0), short (1), tuber- 
1. Pronotum black (0), brown (1), yellow- cle-like (2), absent (3). 
ish (2). 14. Median spine erect (0), prostrate (1). 
2. Abdomen black (0), brown (1), yellow- 15. Median spine stout (0), slender (1). 
ish (2). 16. Antennae densely pubescent (0), not 
3. Hemelytra black (0), brown (1), yel- pubescent (1). 
lowish to white (2), brown with colored 17. Last antennal segment longer than first 
spots (3). and second together (0), shorter than 
4. Body glabrous (0), pilose (1). first and second together (1). 
5. Head convex (0), flat (1). 18. Last antennal segment longer than first 
6. Frontal spines long (0), short (1). (0), shorter than first (1). 
7. Frontal spines erect (0), prostrate (1). 19. Third antennal segment | time as long 
8. Frontal spines stout (QO), slender (1). as the last one (0), 2 times (1), 3 times 
9. Frontal spines in contact or crossed (0), (2);°4 times’ G). 
not in contact (1). 20. Bucculae biseriate (0), triseriate (1), 
10. Occipital spines long (0), short (1). quadriseriate (2), pentaseriate (3). In- 
11. Occipital spines adpressed (0), pros- active character. 
trate (1). The occipital spines are pros- 21. Bucculae open in front (0), closed in 
trate when the spines do not rest onto front (1). 


the head, otherwise they are adpressed. 22. Labium almost extending beyond the 


824 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 3. Matrix of states of the 61 characters for 14 species. 


Taxa Character states 
Nobarnus pilosus 00001100000003?70111012100100?700?701?70004200010010030??202133 
Physatocheila dissimilis 01110100000100001102101110101210?701??0111011111111001??110420 
Cysteochila cubens 2223 00 OVO OOM2 0 Pal 702212 OOMOLOPS 05052703 20 Osa ema TOM 02 O a0 ee 
Parada darlingtoni MOL SOO OOLOOOLOLLLOO2ZINLOPOOLU2112203 21 Ata Os WOM eA Aer OMe 2G 
Parada hackeri Ala aL Sj{0)aL (0) aL (0)(0)(0)(ON0) LaalaE(O)(O)aL{o)sLak(o)tL (0) aa ieeya zat a ee MaE AL ALS} aL gL aL (0) iL ak (OAL aL (0) aL AU (0) al aL al at il (0) 
Parada torta NOM ZO OLA COIN TOO2Z2 TNO LO OL NO 2 VA AAA te eae Oded Ones? AO) @ a2 
Parada absona 0003000111001111100211100001122152110111211111110111110111220 
Parada popla (ONY S4(O)(O) AL TL IL (OV(ON(0) ak iN aL aa) (0) 2A al A S}(0)(0)(0)(O)a al Ast Sy OVA Aral salsa UAL (O)al se (O}A ae ah SL AGO} aN TL AL 3),2,(0) 
Parada solla HL ANS}(0) A110) a) 1 O)(ONCOVaL bea balo(oy Ayal al -A(0)sboj(ojabaal Aral ey zi 2a EAS) aA bs OAL aL LOO) AL aL? 
Parada taeniophora il (0) a1 SOMO AL MAL LOVLONCOVAL wh Oak a7 AA AL Al ALCO) As a a2) A) A aL Sh aA AE ALCO) AL ALCO) 2A ALO) IL WL L200 
Parada majuscula (0) LOS} OO)aL LALO ako) sk Sy aL O\O)IL-7AO) aL (0) ao) a AA) Aa Seay S ya EGE ICSE TE IL (O)IL aL a2) 2) AL aL Lat a 
Parada paitae A ALAL aL (O)(O) AL TL A AL ALG) sb Sy aL (O)(O) ML LAL ICO) (O)(O) aL Aa LAA AL ara GE IETS IL LO) AE AL AL LO) aL IL 12) 2) AL 
Parada minuta APASOO MLL MO MOU PP WO Os SAG OOM UA AO Aakala Wass Lala OAL aL LOO iL i 30 
Parada approba MOA SOMO ssl Pojoy POO) ablOv(o)s} shal sya O)s@ak geal AAs al LNA ALA IL (aL wo) AL AL aL aL LO) AL AL 1. (0) 2,0 


metasternum (0), extending to the mid- 
dle of the metasternum (1), extending 
little beyond the mesosternum (2), 
reaching the posterior margins of the 
mesosternum (3), extending to the mid- 
dle of the mesosternum (4). 


. Mesosternal laminae subparallel (0), 


not subparallel (1). 


. Metasternal laminae wide (Q), narrow 


(1). 


. Sternal laminae open behind (0), closed 


behind (1). 


. Pronotum strongly gibbose (0), slightly 


gibbose (1). 


. Pronotum pubescent (0), glabrous (1). 
. Median carina not areolate (0), uniser- 


iate (1). 


. Median carina with large areolae (0), 


small areolae (1), minute areolae (2). 


. Lateral carinae ridge like (0), with one 


or two rows of areolae (1), with 3 to 5 
rows of areolae (2), with 6 or more 
than 6 rows of areolae (3). Inactive 
character. 


. Lateral carinae erect (0), reflexed but 


not resting onto the pronotum (1), re- 
flexed and resting onto the pronotum 


@): 


. Lateral carinae 8 areolae long (0), 9 


areolae long (1), 10 areolae long (2), 
11 areolae long (3), 12 areolae long (4), 


40. 


41. 


14 areolae long (5), 15 areolae long (6). 
Inactive character. 


. Hood absent (0), present but flat (1), 


cyst-like and partly covering the head 
(2). When the hood is present, it covers 
most of the head but generally not the 
eyes. 


. Hood higher than posterior pronotal 


lobe (0), not higher than posterior pron- 
otal lobe (1). 


. Hood 3 areolae long (0), 4 areolae long 


(1), 5 areolae long (2), 6 areolae long 
(3), 7 areolae long (4). Inactive char- 
acter. 


. Hood 1 areola high (0), 2 areolae high 


(1), 3 areolae high (2). Inactive char- 
acter. 


. Collar triseriate (0), biseriate (1). In- 


active character. 


. Paranota 2 areolae broad (0), 3 areolae 


broad (1), 4 areolae broad (2), 5 areolae 
broad (3). Inactive character. 


. Paranota with large areolae (0), small 


areolae (1), minute areolae (2). 
Paranota more than 14 areolae long (0), 
13-14 areolae long (1), 11—12 areolae 
long (2), 10 areolae long (3), 8—9 are- 
olae long (4). Inactive character. 
Paranota reflexed with free margins in 
contact with pronotum (0), reflexed 
with free margins not in contact with 


VOLUME 102, NUMBER 4 


42. 


43. 


44. 


45. 


46. 


47. 
48. 


49. 
50): 
= 
a2) 


SS: 


54. 


SS) 


56. 
Si 


58. 


50) 


60. 


pronotum (1), not reflexed with free 
margins not in contact with pronotum 
2): 

Paranota not covering pronotum (0), 
covering part of pronotum (1). 
Hemelytra wide (0), narrow (1). Hem- 
elytra are considered narrow when not 
extending far beyond the abdomen. 
Hemelytra sharply widened at base (0), 
not sharply widened at base (1). 
Junction of RM and Cu veins raised 
and forming a little swollen zone (0), 
not raised and forming a little swollen 
zone (1). 

Hemelytra areolae hyaline (0), not hy- 
aline (1). 

Hemelytra pubescent (0), glabrous (1). 
Hemelytral veins raised, forming ridges 
(O), not raised and forming ridges (1). 
Costal area wide (0), narrow (1). 
Costal area plane (0), raised (1). 
Costal area areolae very small (0), 
small (1), large (2), very large (3). 
Costal area 6 areolae broad (0), uni- 
seriate (1). 

Costal area uniseriate at base (0), bi- 
seriate at base along 3-5 areolae (1), 
biseriate at base along 7—10 areolae (2). 
The costal area is mostly uniseriate and 
shows in part two areolae for the same 
width like if the original areolae were 
divided in two smaller areolae (Figs. 
9-15). 

Costal area uniseriate at the middle (0), 
biseriate at the middle along 2-3 are- 
olae (1), biseriate at the middle along 
5—6 areolae (2). 

Subcostal area areolae very small (0), 
small (1), large (2), very large (3). 
Subcostal area large (0), small (1). 
Discoidal area areolae very small (0), 
small (1), large (2), very large (3). 
Discoidal area 4 areolae wide (0), 5 
areolae wide (1), 6 areolae wide (2), 7 
areolae wide (3), 8 areolae wide (4), 9 
areolae wide (5). Inactive character. 
Sutural area areolae very small (0), 
small (1), large (2), very large (3). 
Sutural area 6—7 areolae wide (0), 8—9 


825 


areolae wide (1), 12 areolae wide (2), 
less than 6 areolae wide (3). Inactive 
character. 


The phylogenetic analysis was performed 
with Hennig86 (Farris 1988). The algorithm 
ie* was used to build tree (s), using Fitch 
parsimony. 


RESULTS 


First analysis.—Three trees of 119 
steps, with ci = 61 and ri = 50 were gen- 
erated. They differ by the relative position 
of the three New Caledonian species 
among themselves. Seventy-seven percent 
of the active characters are not informa- 
tive for the resolution of these conflicting 
branches. Here is presented one of the 
trees which corresponds to the following 
interpretation (Fig. 16). For the 7 infor- 
mative characters, Parada minuta and P. 
paitae share the same state for four of 
them, while P. majuscula and P. minuta 
share the same state for one of them, and 
P. majuscula and P. paitae for any of 
them. Then, Parada minuta may be more 
closely related to P. paitae than to P. ma- 
juscula. The coefficients and number of 
steps for each characters is given Table 4. 

The genus Parada (including the species 
approba) is monophyletic. The monophyly 
of the genus is supported by the raised 
veins on the hemelytra, the lateral carinae 
and the paranota which are curved inwards 
but not resting onto the pronotum. 

The genus is divided in two groups, the 
New Caledonian group which comprise 
the three New Caledonian species, and the 
Australian group comprising the other 
eight species. The Australian species have 
five cephalic spines, while New Caledon- 
ian species have four. No unambigous 
synapomorphy support the monophyly of 
the Australian group, while the short oc- 
cipital spine and the lack of a median 
spine are the unambiguous synapomor- 
phies of the New Caledonian group. Par- 
ada approba is completely integrated in 
the genus; however, it is the basal species 


826 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


N. pilosus 


P. dissimilis 


N C. cubens 
56 
P. majuscula 
i P. paitae 
44 Gt) LUG): 
13(3), 19(1) P. minuta 
P. approba 
19(3) 
P. popla 
6(1) 
N P. absona 
314), 37): 35 
41(1), 48(0) P. hackeri 
i 19(1) 
33 5(1) N P. solla 
32 43 
P. darlingtoni 
24(1) 30(2) 
P sorta 
51(2) 
6(1) P. taeniophora 
43 
Fig. 16. One of the three cladograms obtained with the ie* command of Hennig86 on data show in table 3 
(14 taxa, 61 active characters). Length = 119 steps, ci = 61, ri = 50. Paranota and lateral carinae width, 


respectively, in number of areolae are indicated under an arrow on the cladogram. 


of the Australian group. It is characterised have a labial sulcus broad like the New 
by the third antennal segment that is four Caledonian species. Among the Austra- 
times the length of the last. Parada popla, lian group, two subgroups can be distin- 
P. absona and P. approba are the basal guished, the hackeri subgroup composed 
species of the Australian group. They by P. hackeri and P. solla, and the dar- 


VOLUME 102, NUMBER 4 


Table 4. Number of steps, ci and ri for each char- 
acter for the first analysis (10 characters inactive) and 
the second analysis (all characters active). 


ae Analysis 1 Analysis | 
acters Steps ci ri Steps ci ri 

1 5 40 40 5 40 40 

2 5 40 40 5 40 40 

3 5 40 0) 5 40 0) 

4 | 75 O 4 WS 0) 

5 1 100 100 1 100 100 

6 2; 50 66 3 33 33 

7 3) 33 60 3 33 60 

8 2 50 50 2 50 50 

9 2 50 50 2 50 50 
10 2 50 0) 2 50 0) 
11 1 100 100 1 100 100 
12 1 100 100 1 100 100 
13 2 50 50 2 50 50 
14 5) 60 60 5 60 60 
15 2 50 66 3 33 33 
16 1 100 100 1 100 100 
17 1 100 100 1 100 100 
18 1 100 100 1 100 100 
19 i 100 100 1 100 100 
20 4 50 50 4 50 50 
21 >) 40 0) 3) 40 0) 
22 1 100 100 1 100 100 
23) 7 5)// 40 8 50 20 
24 3 33 66 3 33 66 
25 Z 50 TS 4 25 25 
26 3 33 0) 3 33} O 
27 2 50 50 2 50 50 
28 1 100 100 1 100 100 
29 D, 50 0) 2 50 0) 
30 4 50 0) 3 66 50 
31 4 75 O 4 75 0) 
32 1 100 100 1 100 100 
33 U 85 50 7 85 50 
34 3 66 50 2 100 100 
35 2 50 0) 2 50 0) 
36 7 42 0) 6 50 25 
37/ 5 40 0) 3 66 66 
38 1 100 100 1 100 100 
59 5 60 50 3 100 100 
40 2 100 100 2 100 100 
41 7 57 P25) 6 66 50 
42 2 100 100 2 100 100 
43 1 100 100 1 100 100 
44 1 100 100 1 100 100 
45 l 100 100 100 100 
46 4 25 25 3 33 50 
47 1 100 100 1 100 100 
48 1 100 100 1 100 100 
49 i 100 100 1 100 100 
50 1 100 100 1 100 100 
31 1 100 100 1 100 100 


Table 4. Continued. 


Analysis 1 Analysis | 

Char- 

acters Steps ci ri Steps ci ri 
Sy 4 75 Ws 5 60 50 
53 1 100 100 1 100 100 
54 5) 40 25 4 50 50 
55 3 66 10) 3) 66 O 
56 2 100 100 2 100 100 
57 1 100 100 1 100 100 
58 2 100 100 2. 100 100. 
59 9 55 0) 8 62 DS 
60 +4 50 0) 4 50 0) 
61 5 60 33 4 TS 66 


lingtoni subgroup composed by P. dar- 
lingtoni, P. torta and P. taeniophora. The 
monophyly of both subgroups together is 
supported by a narrow labial sulcus. The 
monophyly of hackeri subgroup is sup- 
ported by a flat hood. The monophyly of 
darlingtoni subgroup is supported by very 
large areolae on costal area. The slightly 
gibbose pronotum is autapomorphic for P. 
hackeri, and the large areolae of the me- 
dian carina is autapomorphic for P. torta. 

Second analysis.—A single tree of 168 
steps, with ci = 62 and ri = 48 is gener- 
ated (Fig. 17). The conflicting branch be- 
tween the New Caledonian species is re- 
solved, and Parada minuta is more close- 
ly related to P. paitae than to P. majus- 
cula. As in the first analysis, the genus 
Parada is monophyletic. The monophyly 
being supported by the same characteris- 
tics. Parada hackeri is the basal species 
of the genus. It is the only species of Par- 
ada with stout frontal and occipital 
spines, biseriate bucculae and having the 
shortest rostrum of the genus. It shares the 
same hood shape and the same size of are- 
olae on costal area than Cysteochila cub- 
ens, the outgroup most related to the ge- 
nus. The other species of the genus is di- 
vided in two groups, the torta group, com- 
prising P. torta (basal species of the 
group), P. absona, P. popla, P. darling- 
toni and P. approba, and the solla group, 
comprising P. solla (basal species of the 


828 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


A 
56 
8(1) 
38(1) 
33 
Y 


13@1), 31(1), 19(2) 
STL) ool): 
41(1), 48(0) 


A 
43 38(2) 


6(1), 10(1), 
13(3), 190) & 


N. pilosus 
P. dissimilis 
C. cubens 


P. hackeri 


P. torta 
29(0) 
P. absona 


P. popla 
7(1), 30(2) 
P. darlingtoni 
32 
13(0) P. approba 
19(3) 
P. solla 
S(1) 
P. taeniophora 


P. majuscula 
22(0) 


P. paitae 


P. minuta 


Fig. 17. The single three cladograms obtained with the ie* command of Hennig86 on data show in table 3 
(14 taxa, 51 active characters). Length = 168 steps, ci = 62, ri = 48. Paranota and lateral carinae width, 
respectively, in number of areolae are indicated under an arrow on the cladogram. 


group), P. taeniophora, P. majuscula, P. 
paitae and P. minuta. The torta group is 
characterized by the paranota three areo- 
lae wide, that is shared with Parada hack- 
eri and the outgroup Physatocheila dissi- 


milis. The solla group is characterized by 
the paranota four areolae wide, the only 
autapomorphic character for the group. 
The coefficients and number of steps vary 
for 15 characters over 61 between the two 


VOLUME 102, NUMBER 4 


analysis (Table 4), 6 of them being inac- 
tive in the first one. 


DISCUSSION AND CONCLUSION 


The species of Parada can easily be rec- 
ognized by the combination of the paranota 
(characters 41, 42), the lateral carinae (char- 
acters 30, 31) and the costal area shapes 
(characters 49 to 54). 

Despite the different results of the two 
analyses, some characters have the same 
tendencies of evolution. The lateral carinae 
and the paranota width in terms of number 
of areolae used as attributes in the first ana- 
lysis (because of their uncertain homolo- 
gies) narrow from basal to terminal taxa. 
However, these tendencies in the second 
analysis are not as clear as in the first one. 

In the first analysis, the width of the lat- 
eral carinae narrow from the outgroup C. 
cubens (lateral carinae 6 areolae broad) to 
the New Caledonian group (4 areolae 
broad), the Australian group (3 areolae 
broad) and P. darlingtoni (2 areolae broad), 
except for P. absona which has lateral ca- 
rinae of 5 areolae broad (Figs. 9-15). The 
same way, the paranota narrow from C. 
cubens (5 areolae broad) to the New Cale- 
donian group (4 areolae broad) and the 
Australian group (3 areolae broad). There 
are two reversals for P. solla and P. taenio- 
phora which have the paranota 4 areolae 
broad. The others pronotal and hemelytral 
characters do not show such a tendency. 
However, the lateral carinae and the par- 
anota are longer in terms of number of are- 
olae for C. cubens than for the Parada spe- 
cies, and the lateral carinae of the New Ca- 
ledonian species is shorter than the one for 
the Australian species. 

In the second analysis, the lateral carinae 
narrow from C. cubens to Parada species 
from 6 to 3 areolae wide. They still narrow 
in the torta group for P. darlingtoni from 3 
to 2 areolae wide, while they enlarge in the 
solla group for the three New Caledonian 
species from 3 to 4 areolae. There is also a 
reversion for P. absona which lateral cari- 
nae enlarge to 5 areolae wide. The paranota 


829 


narrow from C. cubens to Parada species 
from 5 to 3 areolae wide but enlarge in the 
solla group to 4 areolae. 

If characters such as the width among the 
different hemelytral areas and the various 
pronotal expansions like the hood, the lat- 
eral carinae and the paranota show a trend 
of evolution, this is not the case of other 
characters such as the length of the lateral 
carinae and the paranota. They do not show 
any extension or shortening through the 
cladogram. Thus, wide lateral carinae and 
wide paranota could be plesiomorphic. The 
two major groups found by the first analy- 
sis, are separated by a geographic barrier: 
the sea between the Queensland and New 
Caledonia, isolated from Australia since the 
Triassic. Then, lateral carinae and paranota 
still narrowed in Australia after the Triassic. 
However, these characters show slight var- 
iation among Parada species, in compari- 
son with other genera. As a general rule, 
Tingidae present traits that have no appar- 
ent biological explanation and that could be 
hypertelic by their disproportionate size. 
But, these possibilities still require further 
study and much more data such as behav- 
ioral parameters, and host plants, need to 
be added to test these evolutionary hypoth- 
eses. 


ACKNOWLEDGMENTS 


I am most grateful to Gordon Nishida (B. 
P. Bishop Museum, Honolulu, Hawaii) for 
the loan of many specimens, and Thomas 
Henry (Systematic Entomology Laboratory, 
USDA) for his helpful support in the access 
to the types at the National Museum of Nat- 
ural History, Smithsonian Institution, 
Washington, DC. I am also indebted to Cyr- 
il d’Haese, Philippe Grandcolas, Laure De- 
sutter-Grandcolas, and Thierry Bourgoin 
(MNHN, Paris) for their fruitful comments. 


LITERATURE CITED 


Drake, C. J., 1942. New Tingidae (Hemiptera). Iowa 
State College Journal of Science 17(1): 1-21. 

. 1952. The genus Parada Horvath (Hemiptera: 

Tingidae). Psyche 59(4): 143-147. 


830 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


. 1961. Some Australian Tingidae (Hemiptera), 
including new genera and new species. Records 
of the Australian Museum 25(6): 107—114. 

Drake, C. J. and FE A. Ruhoff. 1961. New genera and 
new species of lacebugs from the Eastern Hemi- 
sphere (Hemiptera: Tingidae). Procceeding of the 
United States National Museum 113: 125-183. 

. 1965. Lace bugs of the world: a catalog (He- 

miptera: Tingidae). United States National Muse- 

um Bulletin 243, 634 pp. 


Farris, J. S. 1988. Hennig86, version 1.5, Port Jeffer- 
son Station, New York. 

Guilbert, E. In press. New data on New Caledonian 
Tingidae (Hemiptera) with description of new spe- 
cies. Zoologia Neocaledonica Vol. 5. Mémoires 
du Museum National d’Histoires naturelles. 

Horvath, G. 1925. Results of Dr. E. Mj6berg’s swed- 
ish scientific expeditions to Australia 1910— 
1913. 45. Tingitidae. Arkiv for Zoologi 17(24): 
1-17. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 831-837 


A NEW NEOTROPICAL SPECIES OF CLINODIPLOSIS 
(DIPTERA: CECIDOMYIIDAE), AN IMPORTANT NEW PEST OF 
CULTIVATED PEPPERS (CAPSICUM SPP.: SOLANACEAE) 


RAYMOND J. GAGNE, HELGA BLANCO-METZLER, AND JEAN ETIENNE 


(RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. 
Department of Agriculture, % National Museum of Natural History, Smithsonian Insti- 
tution, Washington, DC 20560-0168, U.S.A. (e-mail: rgagne @sel.barc.usda.gov); (HB-M) 
Centro de Investigacion en Proteccién de Cultivos, Estacién Experimental Fabio Baudrit, 
University of Costa Rica, P.O. Box 183-4050, Alajuela, Costa Rica; (JE) Institut National 
de la Recherche Agronomique, Station de Zoologie et Lutte Biologique, Domaine Duclos, 
Petit Bourg, Guadeloupe 


Abstract.—A new species of cecidomyiid, Clinodiplosis capsici Gagné, is reported 
from cultivated sweet and hot pepper in Costa Rica, Guadeloupe, and French Guiana 
where it appears to have potential as a serious pest. Adults, pupae, and larvae are de- 


scribed, illustrated, and compared to other Clinodiplosis species. 


Key Words: 


In the mid-1990s, a new species of gall 
midge belonging to Clinodiplosis was in- 
dependently discovered on Guadeloupe and 
in Costa Rica attacking sweet pepper, Cap- 
sicum annuum L., and hot pepper, Capsi- 
cum frutescens L. The white larvae of the 
gall midge cause swellings on stems, 
leaves, and bases of flowers (Figs. 1-3). 
Galls can be found all year round and, 
where locally abundant, the cecidomyiid is 
a serious pest. Larvae live singly in cells 
within the swellings. The full grown larva 
pupates in the gall, the fully developed 
pupa forms a circular exit hole with the 
help of the antennal horns, emerging about 
halfway out of the hole, and the adult im- 
mediately emerges from the pupal skin. Af- 
ter adult emergence, the damaged part of 
the plant is susceptible to rot. Similar galls 
were later found in 1999 in French Guiana. 

In Costa Rica, galls were collected in dry 
climates as in Guanacaste, wet climates as 
in Pérez Zeled6n, and intermediate cli- 


gall midge, sweet pepper, hot pepper 


mates, such as Alajuela. They were found 
in gardens, backyards and greenhouses of 
Guanacaste, Alajuela, and Cartago Provin- 
ces, but not in commercial plantations. This 
absence in the larger plantations may be 
due to chemical applications intended for 
the control of the pepper weevil, Anthono- 
mus eugenii Caro (Coleoptera: Curculioni- 
dae). 

The new species of gall midge will read- 
ily run to Clinodiplosis in the key in Gagné 
(1994). Clinodiplosis is a worldwide genus 
of some 75 known valid species. Outside 
the Neotropics, most species of this genus 
appear not to be host specific and are usu- 
ally associated with fungus growing in or 
on plant tissue, such as spent flowers or old 
galls. A few species, however, are associ- 
ated with simple bud galls where they are 
doubtless primary plant feeders (Gagné 
1989). While Clinodiplosis spp. may also 
be found associated with fungus in the Neo- 
tropics, most known regional species in this 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 


and closely related genera peculiar to the 
Neotropics, e.g., latrophobia and Schisma- 
todiplosis, are phytophagous, host specific, 
and associated with complex galls (Gagné 
1989), as is the new species described be- 
low. 


METHODS 


Galls with pupae or full-grown larvae 
were placed in small containers until adults 


Galls of Clinodiplosis capsici on stems and leaves of Capsicum sp. 


emerged. Specimens of immature stages 
and reared adults were preserved in 70% 
isopropyl alcohol. Samples were mounted 
on microscope slides using the method out- 
lined in Gagné (1989). Terminology for 
adult morphology follows usage in Mc- 
Alpine et al. (1981) and for larval mor- 
phology that in Gagné (1989). Galls were 
obtained, adults reared, and other stages se- 
cured by H. Blanco-Metzler in Costa Rica 


VOLUME 102, NUMBER 4 


and by J. Etienne in Guadeloupe. J. Etienne 
also found similar galls in French Guiana. 
The taxonomic investigation was the re- 
sponsibility of R. J. Gagné. 


Clinodiplosis capsici Gagné, new species 
(Figs. 4—15) 


Adult.—Head: Eyes connate, 11—12 fa- 
cets long at vertex; facets mostly hexago- 
noid, all closely adjacent. Occiput with dor- 
sal protuberance with 2 apical setae. Frons 
with 6-10 setae. Labella ellipsoid and 
pointed apically, each with several lateral 
setae. Palpus 4-segmented. Male antennal 
flagellomeres (Fig. 4) binodal; one circum- 
filum on basal node, two on the distal, the 
loops of the three circumfila subequal in 
length. Female flagellomeres (Fig. 5) cylin- 
drical with long necks, surrounded by two 
appressed circumfila connected by two lon- 
gitudinal bands. 

Thorax: Wing unmarked, 2.2 to 2.9 mm 
long, R; curved toward apex, joining C pos- 
terior to wing apex. Tarsal claws (Fig. 6) 
untoothed, curved near basal third; empodia 
very short, not attaining bend in claws. 

Male abdomen: First through sixth ter- 
gites entire, rectangular, with single poster- 
ior row of setae, several lateral setae, scat- 
tered scales, and 2 anterior trichoid sensilla; 
seventh tergite unsclerotized posteriorly 
and lacking the posterior row of setae and 
scales, but lateral setae and anterior pair of 
trichoid sensilla present; eighth tergite un- 
differentiated, the only vestiture the anterior 
pair of trichoid sensilla. First through eighth 
sternites rectangular, covered with setae and 
with 2 anterior trichoid sensilla; eighth ster- 
nite similar to preceding except weakly 
sclerotized anterolaterally. Genitalia (Figs. 
9-10): cerci rectangular, with posterior se- 
tae; hypoproct much longer than cerci, wid- 
est at midlength, narrowing beyond mid- 
length and widening again posteriorly, the 
basal two-thirds expanded laterally to curve 
slightly around aedeagus, and with strong 
setae and several smaller ones posterolat- 
erally; aedeagus elongate, narrower and 
longer than hypoproct, with longitudinal 


833 


rows of sensory pits; gonocoxite elongate- 
cylindrical with mesoposterior surface 
forming nearly right angle; gonostylus 
elongate-cylindrical, with setulae near base 
and covered beyond with minute carinae 
and widely scattered short setae. 

Female abdomen (Figs. 7-8): First 
through seventh tergites entire, rectangular, 
with mostly single row of posterior setae, 
several lateral setae, extensively covered 
with scales, and with 2 anterior trichoid 
sensilla. Eighth tergite unsclerotized, with 
mostly single row of posterior setae and an- 
terior pair of trichoid sensilla the only ves- 
titure. Second through seventh sternites 
quadrate, extensively covered with setae 
and scales and with anterior pair of trichoid 
sensilla. Ovipositor slightly protrusible, 
venter of eighth segment and dorsum of 
ninth and tenth segments without vestiture, 
venter of ninth and tenth segments with se- 
tae, cercus large, ovoid, with pair of apical 
sensory setae and scattered setae elsewhere, 
hypoproct short, narrow, with 2 posterior 
setae. 

Pupa.—Head (Figs. 11-12): Antennal 
base pointed apically, the apex projecting 
ventrad; cervical sclerite with two elongate 
setae; face without ventral projections, with 
1—2 papillae, one with seta, on each side of 
base of labrum. Prothoracic spiracle elon- 
gate, pointed apically. Abdominal tergites 
covered dorsally with uniformly small spic- 
ules, none enlarged and spiniform. 

Third larval instar (Figs. 13—15).— 
Length, 2.5—2.7 mm. White. Integument 
with scattered spicules. Antenna about 
twice as long as wide. Spatula with 2 wide- 
ly separated anterior teeth, the space be- 
tween weakly indented. Lateral thoracic pa- 
pillae in 2 groups of 3 on each side of cen- 
tral line, 2 papillae in each group each with 
tiny seta. Dorsal and pleural papillae elon- 
gate. Terminal segment rounded, with 8 pa- 
pillae as follows: 1 pair as long as dorsal 
setae of previous segment; 1 pair with setae 
about half as long; the two posterior pairs 
with short, corniform setae, those of inner 
pair slightly narrower than outer pair. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


‘@ 


Figs. 2-3. 2, Galls of Clinodiplosis capsici on stems and base of flowers of Capsicum sp. 3, Detail of same, 


one in section to show larva in larval cell. 


VOLUME 102, NUMBER 4 835 


? 
\ 
\ 
\ 
‘ | 
XX 

< 10 

Figs. 4-10. Clinodiplosis capsici. 4, Male third antennal flagellomere. 5, Female third antennal flagellomere. 
6, Tarsal claws and empodium. 7, Female postabdomen, seventh segment to cerci (lateral). 8, Female cercus, 


detail (lateral). 9, Male genitalia, only one gonopod shown (dorsal). 10, Same, only one gonocoxite shown 
(ventral). 


836 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 11-15. Clinodiplosis capsici. 11, Pupal head (ventral). 12, Pupal head, part, showing ventral projection 
of antennal base (lateral). 13, Larval spatula and associated lateral papillae. 14, Larva (ventral). 15, Larval eighth 
and terminal segments (dorsal). 


VOLUME 102, NUMBER 4 


Holotype.—¢d, from Capsicum frutes- 
cens, Guadeloupe, French Antilles, Ravine 
Chaude, 27-XII-1994, J. Etienne, GR 1575, 
deposited in the National Museum of Nat- 
ural History (USNM), Washington, DC. 

Other material examined (all deposited in 
USNM).—Same data as holotype, 1 d6, 2 
2, 3 pupae, 3 larvae; from Capsicum sp., 
Guadeloupe, Ste. Rose, 3-II-1993, J. Etien- 
ne, GR 1309, 2 6, 2 2, 5 pupae, 6 larvae; 
from Capsicum sp., Guadeloupe, Lamentin, 
1-VII-1992, J. Etienne, GR 1110, 1 d, 1 2, 
1 pupa, | larva; from Capsicum annuum, 
Costa Rica, Guanacaste Province, 3-I-1995, 
H. Blanco, 1 3d, 1 @; from Capsicum an- 
nuum, Costa Rica, Turialba, VIII-1995, H. 
Blanco; 36; 22. 3 pupae. 

Etymology.—The specific name, capsici, 
means “of capsicum.” 

Remarks.—The larva of C. capsici dif- 
fers substantially from other Clinodiplosis 
spp. in that the corniform setae of the ter- 
minal segment are small and not each sit- 
uated at the end of lobes, as is usual for the 
genus. It cannot be said with confidence 
whether in this species the lack of lobes is 
primitive or the result of reduction. The 
male and female of C. capsici have the gen- 
eral habitus of the genus, the only pecu- 
liarity being the shape of the male hypo- 
proct, which is broadened at midlength, its 
sides bending slightly around the aedeagus. 
The pupal abdomen is without dorsal spines 
and resembles in that way the species of 
Clinodiplosis that feed on fungi. Unlike the 


837 


new species, several other Neotropical spe- 
cies of Clinodiplosis that pupate in their 
galls, e.g., Clinodiplosis eupatorii Felt in 
conical leaf galls on Chromolaena (Aster- 
aceae) (Gagné 1977), have enlarged dorsal 
spines on the pupal abdomen that are pre- 
sumably adaptations in those species for es- 
caping from the galls. 


ACKNOWLEDGMENTS 


We thank P. Malikul for making the slide 
preparations, Lucrecia Rodriguez for com- 
puter assistance in making the plates, and, 
for their comments on drafts of the manu- 
script: P. Hanson, Universidad de Costa 
Rica, San José; K.M. Harris, International 
Institute of Entomology, CAB, London, 
UK; P. Kolesik, The University of Adelaide, 
SA, Australia; and A.S. Konstantinov and 
A.L. Norrbom of the Systematic Entomol- 
ogy Laboratory, USDA, Washington, DC. 


LITERATURE CITED 


Gagné, R. J. 1977. The Cecidomyiidae (Diptera) as- 
sociated with Chromolaena odorata (L) K. & R. 
(Compositae) in the Neotropical Region. Brenesia 
12/13: 113-131. 

1989. The Plant-Feeding Gall Midges of 

North America. Cornell University Press, Ithaca, 

New York, xiii & 355 pp. & 4 pls. 

. 1994. The Gall Midges of the Neotropical Re- 
gion. Cornell University Press, Ithaca, New York, 
xv & 352 pp. 

McAlpine, J. E, B. V. Peterson, G. E. Shewell, H. J. 
Teskey, J. R. Vockeroth, and D. M. Wood, eds. 
1981. Manual of Nearctic Diptera, Vol. 1. Re- 
search Branch, Agriculture Canada, Monograph 
27, vi & 674 pp. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 838-842 


NEW SPECIES OF PUERTO RICAN PHYCITINAE 
(LEPIDOPTERA: PYRALIDAE) 


H. H. NEUNZIG 


Department of Entomology, North Carolina State University, Raleigh, NC 27695-7613, 


U.S.A. 


Abstract.—Zamagiria borinquensis and Oryctometopia maricaoensis are described as 
new species. Both were collected as adults in western Puerto Rico. 


Key Words: 


Post-Columbian Puerto Rico has been 
subjected to major environmental changes. 
Agriculture, mainly the extensive growing 
for many years of sugarcane and coffee, re- 
placed much of the native vegetation. Some 
refugia exist, however, chiefly as parks or 
reserves. These vary from tropical dry for- 
est to tropical rain forests. Recently, ento- 
mologists at the Carnegie Museum of Nat- 
ural History (CMNH) have intensively 
light-trapped Lepidoptera in these remain- 
ing pockets of relatively undisturbed flora. 
Two previously unknown species of phy- 
citines were collected. These are described 
below, and are added to the 20 species (Ta- 
ble 1) previously recorded from Puerto 
Rico by Heinrich (1956). 


Zamagiria borinquensis Neunzig, 
new species 
(Figs. 1, 3-5) 


Type locality.—Puerto Rico, Isabela, 
Bosque Estatal de Guajataca, Montanas Ay- 
mamon. 

Diagnosis.—The male and female geni- 
talia can be used to separate Zamagiria bor- 
inquensis from other species in the genus. 
Features peculiar to the new species are the 
short, triangular inner clasper on the inner 
base of the valva, the somewhat elongate, 
spoonshaped apical gnathal process of the 


moths, phycitines, Puerto Rico 


male (Fig. 3), and the girdle of robust 
spines in the corpus bursae of the female 
(Fig. 5). 

Description.—Forewing length 6.5—7.0 
mm. Head brownish white. Labial palpus 
white and fuscous. Maxillary palpus white 
to brownish white, aigrettelike in male. 
Dorsum of thorax brownish white suffused 
with fuscous. Forewing brown dusted with 
white; black ridge of raised, curled scales 
preceding antemedial line; antemedial line 
white, weakly developed, with associated 
adjacent bands of dark scales; postmedial 
line similar, in color and development, to 
antemedial line; discal spots black, moder- 
ately distinct, separate; patch of very pale 
reddish brown scales at inner base of wing 
and scattered patches of darker reddish 
brown on outer % of wing. Underside of 
male fore and hindwing without patches or 
streaks of contrasting colored scales (fre- 
quently seen in other Zamagiria). Male 
genitalia (Figs. 3, 4): Uncus triangular, nar- 
rowly rounded distally; apical process of 
gnathos expanded into a somewhat elongate 
spoon-shaped process; juxta a U-shaped, 
thin plate with setiferous lateral arms; tran- 
stilla absent; valva slender, unarmed except 
for triangular, erect clasper on its inner 
base; aedoeagus with single, short, stout, 
well-sclerotized cornutus; vinculum about 


VOLUME 102, NUMBER 4 


Table 1. Phycitinae listed by Heinrich (1956) as 
occurring in Puerto Rico!. 


Crocidomera fissuralis (Walker) 
Ectomyelois furvidorsella (Ragonot) 
Fundella ignobilis Heinrich 
Difundella distractor Heinrich 
Anadelosemia texanella (Hulst) 
Davara interjecta Heinrich 
Davara rufulella (Ragonot) 
Sarasota furculella (Dyar) 
Caristanius pellucidellus (Ragonot) 
Eurythmasis ignifatua Dyar 
Paconius corniculatus Heinrich 
Mescinia berosa Dyar 

Phestinia costella Hampson 
Laetilia portoricensis Dyar 
Metephestia simplicula (Zeller) 
Moodnopsis portoricensis Heinrich 
Prosoeuzophera impletella (Zeller) 
Caudellia clara Heinrich 

Ribua contigua Heinrich 

Varneria dubia Heinrich 


! List does not include a few widespread neotropical 
phycitines that Heinrich included in his checklist as 
occuring in his category “‘West Indies,’ some of 
which may be found in Puerto Rico. 


as long as greatest width. Female genitalia 
(Fig. 5): Ductus bursae short, about / as 
long as corpus bursae, narrow, except 
slightly broadened near ostium bursae, 
without sclerotized plates common to most 
other Zamagiria species, but generally 
sclerotized anteriorly; corpus bursae elon- 
gate, with well-developed, irregular comb 
of many slender spines extending distally 
from ductus bursae to about midway on 
corpus, and with strongly formed band en- 
circling corpus bursae at about middle and 
bearing about 24 robust spines; ductus sem- 
inalis attached to corpus bursae near junc- 
tion of ductus bursae and corpus bursae. 
Type material—Holotype: ¢: Puerto Rico, 
Isabela, Bosque Estatal de Guajataca, Mon- 
tanas Aymamon, 18°25'06N, 66°57'55’W, 
forest, 210 m, 14—15 June 1996, J. Rawlins, 
W. Zanol, R. Davidson, C. Young, M. Klin- 
gler, S. Thompson; genitalia slide HHN 
4427. Paratype: 1 2, same collection data 
as for holotype; genitalia slide HHN 4428. 


839 


Jae, 15 2. 
lotype. 2, Oryctometopia maricaoensis, holotype. 


Males. 1, Zamagiria borinquensis, ho- 


Holotype and paratype deposited in 
CMNH. 

Remarks.—Zamagiria borinquensis is 
very similar externally to other species in 
the genus. The character states of the male, 
particularly the grooved head with its un- 
usual compact tuft of scales attached to the 
frons, and the unique tuft, composed of sev- 
eral distinct types of scales, at the base of 
the antenna, are easily recognized traits of 
the species. 

Etymology.—The specific epithet is de- 
rived from the Taino name for Puerto Rico 
(Borinquen), and the Latin suffix -ensis (de- 
noting place). 


Oryctometopia maricaoensis Neunzig, 
new species 
(Figs. 2, 6-8) 


Type locality.—Puerto Rico, 3.3 km. sw 
Maricao, Bosque Estatal de Maricao. 


840 


Figs. 3—5. 
bursae and corpus bursae. 


Diagnosis.—The male is readily identi- 
fied by the elongate, medially constricted 
vinculum (Fig. 6), and the female by the 
large spined plate that covers most of one 
side of the corpus bursae (Fig. 8). 

Description.—Forewing length 10.5— 
12.0 mm. Head of male brownish white to 
pale reddish brown suffused with reddish 
brown, and with vertex sulcate and white; 
upper part of frons produced dorsally into 
a ridge. Head of female simple, brownish 
white to pale reddish brown. Labial palpus 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Zamagiria borinquensis. 3, Male genitalia (aedoeagus omitted). 4, Aedoeagus. 5, Female ductus 


brown dusted with white and with a few 
scattered brownish red scales. Maxillary 
palpus of male aigrettelike, mostly pale 
brown, in part pink. Maxillary palpus of fe- 
male simple, pale brownish white. Dorsum 
of thorax pale brown or pale purplish 
brown. Forewing pale purplish brown in 
basal 4%; antemedial line moderately distinct 
to distinct, white, its inner % preceded by 
black and pale reddish brown patch, and 
followed by black line and associated white 
patch; postmedial line moderately distinct 


VOLUME 102, NUMBER 4 


Figs. 6-8. 
ductus bursae, corpus bursae and anterior part of ductus seminalis. 


to distinct, white; medial area with patches 
of red to dark red mostly along costa and 
at discal spot; subterminal area with broad 
brownish red patch following postmedial 
line. Male genitalia (Figs. 6, 7): Uncus nar- 
rowly triangular, rounded at apex; apical 
process of gnathos a well-developed hook; 
juxta platelike with thin, inwardly curved 
lateral arms; transtilla absent; valva simple, 
with short, stout lobes near inner base; ae- 
doeagus, long, slender; vesica with cluster 


841 


Oryctometopia maricaoensis. 6, Male genitalia (aedoeagus omitted). 7, Aedoeagus. 8, Female 


of small spines; vinculum long and slender 
(about 2% times as long as basal width), and 
constricted medially. Female genitalia (Fig. 
8): Ductus bursae slightly shorter than cor- 
pus bursae, sclerotized over % its length; 
corpus bursae with large, many-spined 
plate, covering most of left side (when 
viewed ventrally) of bursae; plate, in part 
perforate; ductus seminalis attached to cor- 
pus bursae near junction of ductus bursae 
and corpus bursae. 


842 


Type material.—Holotype: d¢: Puerto 
Rico, Bosque Estatal de Maricao, 3.3 km 
sw Maricao, 18°09'39"N, 67°00'05’W, for- 
est, 550 m, 10-11 June 1996, J. Rawlins, 
C. Young, R. Davidson, W. Zanol, S. 
Thompson, M. Klingler; genitalia slide 
HHN 4459. Paratypes: (4 ¢, 2 @, total); 
same collection data as for holotype, geni- 
talia slides HHN 4415, 4416 (3 6, 2 2); 
Puerto Rico, Isabela, Bosque Estatal de Gua- 
jataca, Montanas Aymamon, 18°25'06’N, 
66°57'55"W, forest, 210 m, 14-15 June 
1996, J. Rawlins, W. Zanol, R. Davidson, 
C. Young, M. Klingler, S. Thompson (1 ¢). 
All specimens from Maricao deposited in 
CMNH,; Isabela paratype in North Carolina 
State University Collection (NCSU). 

Remarks.—The new species differs most 
noticeably from Oryctometopia fossulatella 
Ragonot, the only other species in the ge- 
nus, in that the male genitalia lack the short, 
thornlike spines projecting from the lower 
margin of the sacculus, and have a cluster 
of small spines on the vesica (rather than a 
single spinelike cornutus), and in that the 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


female ductus bursae is more extensively 
sclerotized. Nevertheless, males of both 
species have an enlarged frons and sunken 
vertex, tufted and otherwise modified an- 
tennae, simple scale tufts on abdominal seg- 
ment VIII, and generally similar genitalia. 
Females of both have a characteristically 
large, sclerotized plate in the corpus bursae 
bearing numerous short spines. 
Etymology.—tThe specific epithet is de- 
rived from the locality (Maricao) where al- 
most all the specimens were collected, and 
the Latin suffix- ensis (denoting place). 


ACKNOWLEDGMENTS 


J. E. Rawlins generously made available 
specimens for study, R. L. Blinn photo- 
graphed the new species, and L. L. Deitz, 
D. L. Stephan, and B. M. Wiegmann re- 
viewed an initial draft of the manuscript. 


LITERATURE CITED 


Heinrich, C. 1956. American moths of the subfamily 
Phycitinae. United States National Museum Bul- 
letin 207: 1-581. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 843-851 


TAXONOMY AND POLYTENE CHROMOSOMES OF 
SIMULIUM PARNASSUM MALLOCH (DIPTERA: SIMULIIDAE) 


Eric S. PAYSEN AND PETER H. ADLER 


Department of Entomology, Clemson University, Clemson, SC 29634-0365, U.S.A. 
(e-mail: padler@clemson.edu) 


Abstract.—The taxonomic status of Simulium parnassum Malloch was investigated cy- 
tologically and morphologically. Specimens were collected at 44 localities from the Gaspé 
Peninsula of Quebec to northern Alabama. A previously unreported pupal form of S. 
parnassum from two streams in South Carolina and one stream each in North Carolina 
and Massachusetts lacked the typical rugose sculpturing on the dorsum of the head and 
thorax. No additional structural characters in any life stage differentiated the two forms. 
A standard map of the silk-gland chromosomes was constructed and compared with the 
standard map for the subgenus Simulium. Simulium parnassum differed from the subge- 
neric standard by six inversions in IL, one in IIL, three in IIS, and an unresolved number 
in IL. Placement of S. parnassum in a separate species group, the S. parnassum species 
group, in the subgenus Simulium is suggested on the basis of cytological evidence. No 
fixed rearrangements were found among populations of S. parnassum from Quebec to 
Alabama nor between populations of smooth and rugose pupal forms. A Y-linked poly- 
morphism was observed in the centromere region of chromosome I in some males, but it 
was not associated with either pupal form. Present evidence indicates that S. parnassum 


is a single species throughout its range. 


Key Words: 


Simulium parnassum is one of the most 
distinctive members of the subgenus Si- 
mulium in the Nearctic Region (Stone 
1964). The female is shiny black and each 
claw is long and nearly sigmoidal, with a 
small tooth near its midlength. The pupal 
gill of six filaments and the simple, slipper- 
shaped cocoon are similar to those of the S. 
tuberosum and S. venustum species groups, 
but the rugose head and thorax are unique. 
The deeply incised, triangular postgenal 
cleft of the larva is distinctive. Despite the 
unique attributes of S. parnassum, this spe- 
cies has been assigned to the S. tuberosum 
species group (Crosskey and Howard 
1997), ostensibly because of perceived sim- 
ilarities in the male genitalia, notably the 


Simulium parnassum, black flies, aquatic insects, polytene chromosomes 


presence of a medially directed lobe on the 
gonostylus. The homology of this lobe, 
however, is questionable based on differ- 
ences in location, size, and vestiture. Ac- 
cordingly, a reevaluation of the phyloge- 
netic placement of S. parnassum is war- 
ranted. 

Simulium parnassum was described from 
females taken in northern New Hampshire 
(Malloch 1914). Dyar and Shannon (1927) 
described a male from eastern Virginia as 
S. hydationis, which was synonymized with 
S. parnassum by Stone and Jamnback 
(1955). All life stages of S. parnassum have 
been described and figured (Stone and 
Jamnback 1955, Davies and Peterson 1956, 
Davies et al. 1962, Wood et al. 1963, Stone 


844 


Table 1. Numbers of two pupal forms (rugose, 
smooth) of Simulium parnassum collected at Smith 
and Abner Creeks in Pickens County, South Carolina, 
1998-1999. 


Smith Creek Abner Creek 
Ru- Ru 
Year Date gose Smooth Date gose Smooth 
1998 25 April D2 0) 2 June Ope 
1998 5 May 0) + 9 June O 20 
1998 14 May 7) 10) 11 June 1 23 
1998 23 May! 4 23 June! 0) ] 
1999 10 May ] 0) 1 June 1 12 
1999 15 May 1 1 5 June Orel 
1999 18 May 1 5) 10 June! QO 26 
1999 21 May 30-36 
1999 24 May 1 30 
1999 28 May 0 17 
1999 1 June! 0) 4 


' No pupae found beyond these dates. 


and Snoddy 1969, Adler and Kim 1986). 
The polytene chromosomes have not been 
studied, although Rothfels (1979) com- 
mented briefly on the banding sequence of 
two of the six chromosomal arms. Conse- 
quently, S. parnassum has not been inves- 
tigated for sibling species, which are com- 
mon among black flies (Adler 1988). 

This species occurs in eastern North 
America from Canada to the southern end 
of the Appalachian Mountains (Stone and 
Snoddy 1969), with an isolated record from 
Missouri (Doisey et al. 1986). The imma- 
ture stages occupy cool, rocky, forest 
streams (Davies et al. 1962, Adler and Kim 
1986). Females are mammalophilic (Fuller 
1940, Downe and Morrison 1957, Addison 
1980) and can be pests of humans (Adler 
and Kim 1986, Gibbs et al. 1986). 

Our objectives were to screen S. parnas- 
sum for sibling species, using morphology 
and polytene chromosomes, and to resolve 
its chromosomal banding sequence relative 
to the subgeneric standard of Rothfels et al. 
(1978), with the intent of gaining phylo- 
genetic insight. 


MATERIALS AND METHODS 


Larvae and pupae of S. parnassum were 
collected from 1998 to 1999 at 44 sites 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


from the Gaspé Peninsula of Quebec to 
northern Alabama, and were fixed in Car- 
noy’s solution (1 part glacial acetic acid: 3 
parts 95% ethanol). Additional pupae were 
reared individually to adults on moist filter 
paper in petri dishes. 

Seasonal sampling of larvae and pupae 
was conducted at Smith Creek (35°0.17'N 
82°49.01'W) and Abner Creek (35°4.27'N 
82°47.11'W) in Pickens Co., South Caroli- 
na. Streams were sampled every 10 days in 
1998 and every three days in 1999. In 1998, 
Smith Creek was sampled 3 March—11 Au- 
gust and Abner Creek 13 May—11 August. 
In 1999, both streams were sampled 17 
March—26 June. 

Polytene chromosomes were prepared 
from larval silk glands, using the Feulgen 
method of Rothfels and Dunbar (1953). A 
standard chromosome map was constructed 
for S. parnassum, following the procedures 
and nomenclature of Rothfels et al. (1978). 
The banding pattern of the standard map 
was compared with that of the subgeneric 
standard of Rothfels et al. (1978). For chro- 
mosomal arms of S. parnassum that had 
banding patterns identical to those of the 
subgeneric standard, the subgeneric num- 
bering was preserved. Chromosome arms 
of S. parnassum that were rearranged rela- 
tive to the subgeneric standard were given 
new section numbers to provide a continu- 
ous sequence. Slide-mounted chromosomes 
of larvae from 15 of the 44 collection sites 
(see Chromosomal Material Examined) 
were compared band for band against the 
standard map of S. parnassum in a search 
for rearrangements within and between 
populations of S. parnassum. Specimens 
and chromosomal photographic negatives 
and working maps were deposited in the 
Clemson University Arthropod Collection, 
Clemson, South Carolina. 

Chromosomal material examined (num- 
bers of each sex refer to larvae for which all 
chromosomal bands were read).—ALA- 
BAMA: Talladega Co., Cheaha State Park, 
Cheaha Creek, 33°28.27'N 85°49.07'W, 27 
March 1998 (1 3); 19 May 1998 (9 @); 


VOLUME 102, NUMBER 4 845 


Figs. 1-2. Cephalic plates of pupal exuviae of Simulium parnassum. 1, Rugose form (Smith Creek, Pickens 
Co., South Carolina, 18 May 1999). 2, Smooth form (Abner Creek, Pickens Co., South Carolina, 11 June 1998). 


846 


pe 


Figs. 3-5. 


3 3) 
ie! > \ >, 
4] all 
al 34 32 | - v35\32\ > 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Silk-gland chromosomes of Similium parnassum, with landmarks of Rothfels et al. (1978); C = 


centromere region. 3, Chromosome arm IS. Numbers indicate section numbers for both S. parnassum standard 
sequence and Simulium subgeneric standard of Rothfels et al. (1978). Male from South Carolina, Pickens Co., 
Oil Camp Creek, 8 May 1998 (sections 1 to center of section 13) plus female from Massachusetts, Berkshire 
Co., 29 June 1998 (section 20 to center of section 13); Gl = glazed, 2 = two blocks, 3 = three heavy. 4, 
Chromosome arm IL. Numbers on top of chromosome indicate S. parnassum standard banding sequence; bottom 
numbers correspond with those of Simulium subgeneric standard of Rothfels et al. (1978). Brackets indicate 
inversions relative to Simulium subgeneric standard of Rothfels et al. (1978); numbered brackets indicate se- 
quence in which overlapping inversions occurred. Arrows indicate breakpoints for complex of three inversions 
relative to subgeneric standard. Female from Georgia, Dade Co., 19 May 1998; Ma = marker, Sp = spongy. 5, 
Centromere region of chromosome I. Male from Georgia, Dade Co., 19 May 1998; C’ = condensed, Y-linked 


centromere band. 


Cheaha State Park, Dry Creek, 33°28.23'N 
85°48.72'N, 19 May 1998 (1 6, 6 Q); 
GEORGIA: Dade Co, Johnson’s Crook, 
Newsome Gap Road, 0.8 km west of Moore 
Rd., 34°47.23'N 85°28.25'W, 19 May 1998 
(2 6,7 2); MASSACHUSETTS: Berkshire 
Co., Jug End Rd., 0.24 km west of Guilded 
Hollow Road, 42°8.99'N 73°26.98'W, 29 
June 1998 (2 36, 16 2); NEW HAMP- 
SHIRE: Carroll Co., Bear Notch Rd., 3.2 
km “north “ef “SR 112." “44° 1750N 
71°19.13'W, 24 July 1998 (1 3); Grafton 
Co., SR118, 10.5 km southwest of SR 112, 


43°59.00'N 71°47.93'W, 24 July 1998 (2 6, 
7 2); NEW JERSEY: Sussex Co., Tillman 
Brook, 41°0.15'N 74°0.86'W, 22 June 1998 
(1 36, 4 2); NORTH CAROLINA: Hay- 
wood Co., US Rt. 276, 1.6 km north of Cru- 
so, 35°26.4'N 82°48.8’W, 7 June 1998 (2 
6); Madison Co., Silver Mine Creek 
35°0.53'N 82°0.48'W, 14 May 1998 (3 gd, 
2 2); SOUTH CAROLINA: Pickens Co., 
Abner Creek, SR 1105, 4.8 km east of Hwy 
178, 35°4.27'N 82°47.11W’, 23 May 1998 
(14 6,5 2); 2 June 1998 (10 2); Oil Camp 
Creek, Oil Camp Creek Rd., 0.8 km west 


VOLUME 102, NUMBER 4 


847 


Fig. 6. Chromosome arm IIS of Simulium parnassum, with landmarks of Rothfels et al. (1978). C = cen- 
tromere region. Section numbers on bottom indicate S. parnassum standard banding sequence; those on top refer 
to Simulium subgeneric standard sequence of Rothfels et al. (1978). Bracket indicates B inversion of Rothfels 
et al. (1978). Arrows indicate breakpoints of two overlapping inversions relative to subgeneric standard. Female 


from Massachusetts, Berkshire Co., 29 June 1998. Bu = bulge, Rb = ring of Balbiani, Tr = trapezoidal. 


of River Falls Rd., 35°6.65'’N 82°34.08’W, 
8 May 1998 (7 6, 1 2); TENNESSEE: 
Monroe Co., Buckhorn Cr., 35°19.98'’N 
84°9.83'W, 6 June 1998 (2 3); VIRGINIA: 
Augusta Co., East Dry Branch, SR 688, 3.2 
km: north of SR 42, 38°12.70'N 
79°16.31'W, 25 June 1998 (8 2); NEW 
BRUNSWICK: Restigouch Co., Collector 
Hwy. 180, 37 km west of Bathurst, 27 July 
1998 (3 3, 12 2); QUEBEC, Gaspé Pen- 
insula, Hwy. 132, 5.6 km south of Routh- 
ierville, 29 July 1998 (5 6, 19 @). 


RESULTS 


Two pupal forms of S. parnassum were 
discovered among the 472 pupae examined. 


Rugose pupae had a raised pattern of retic- 
ulation on the dorsal surface of the head 
and thorax (Fig. 1). Smooth pupae lacked 
surface sculpturing, although the pattern of 
rugosity could be seen, with substage light- 
ing, as a vague outline on the head and tho- 
rax of the pupal exuviae (Fig. 2). Interme- 
diates between the smooth and rugose 
forms were not found. No additional mor- 
phological characters in larvae, pupae, or 
adults were found that correlated with the 
smooth and rugose pupal forms. 

Smooth pupae were collected from four 
streams. They were found with the rugose 
form in streams in Pickens Co., South Car- 


Fig. 7. 


Chromosome arm IIIS of Simulium parnassum, with landmarks of Rothfels et al. (1978). C = 


centromere region. Section numbers are those of both S. parnassum standard banding sequence and Simulium 
subgeneric standard of Rothfels et al. (1978). Male from South Carolina, Pickens Co., Oil Camp Creek, 8 May 
1998. Bl = blister, Ca = capsule. 


848 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 8-9. Silk-gland chromosomes of S. parnassum, with landmarks of Rothfels et al. (1978). C = centro- 
mere region. 8, Chromosome arm IIL. Section numbers refer to Simulium parnassum standard banding sequence. 
Bracket indicates simple inversion relative to Simulium subgeneric standard of Rothfels et al. (1978). Female 
from Tennessee, Monroe Co., 6 June 1998 (sections 55 through 70) plus female from North Carolina, Haywood 
Co., 7 June 1998 (section 71). DNA = DNA puff, gB = gray band, Po = polar, St = saw tooth, Sy = 


VOLUME 102, NUMBER 4 


olina (Smith and Abner creeks), where they 
were the predominant form, and in Berk- 
shire Co., Massachusetts. In Transylvania 
Co., North Carolina (South Prong Gladdy 
Fork), one smooth pupa and no rugose pu- 
pae were collected. Pupae from near the 
type localities of S. parnassum and S. hy- 
dationis were rugose. No differences in sea- 
sonality were detected in streams where 
both pupal forms occurred (Table 1), al- 
though the rugose form was infrequent in 
these streams. Seasonal differences, how- 
ever, might have been apparent if popula- 
tions of each form had been larger. 

In Smith Creek, where a minimum-max- 
imum thermometer had been placed, larvae 
first appeared on 13 March 1998; water 
temperature was 3.0—9.0°C for the three 
days prior to collection. Pupae were first 
found in Smith Creek on 25 April 1998; 
water temperature was 11.0—12.0°C during 
the 11 days prior to collection. In 1998, the 
last larvae were found at Smith Creek on 2 
June (water temperature for 9 days prior to 
collection = 17.5—18.5°C) and in 1999 on 
1 June (water temperature for 3 days prior 
to collection = 17.5—18.0°C). Pupae were 
first collected from Abner Creek on 2 June 
1998 (temperature at time of collection = 
18.0°C) and 1 June 1999 (16.0°C). The last 
larvae and pupae were collected from Ab- 
ner Creek on 23 June 1998 (18.0°C) and 10 
Jone 19995(15:0°C), 

Of 211 larvae prepared for chromosomal 
analysis, 72% were read, band for band, in 
their entirety. The banding sequences of the 
short arms of chromosomes I and III (Figs. 
3, 7) matched those of the subgeneric stan- 
dard, although we were unable to match all 
of the fine bands in the base of IS (sections 
19 and 20). The remaining four arms had 
fixed rearrangements relative to the subge- 


849 


neric standard. IL had one central inversion, 
two overlapping basal inversions, and a 
complex of three inversions distally (Fig. 
4). ILS carried the B inversion of Rothfels 
et al. (1978), plus two additional inversions 
(Fig. 6). IfL had one central inversion (Fig. 
8). The ITIL arm, with the nucleolar orga- 
nizer displaced more distally relative to the 
subgeneric standard, was highly rearranged 
(Fig. 9). Many bands could be recognized, 
but we conservatively matched only the 
bands in the base of IIIL, relative to the 
subgeneric standard, and indicated the char- 
acteristic “‘marker’’? (sensu Rothfels et al. 
1978), which was partitioned by rearrange- 
ments into two pieces. 

We found no fixed or floating inversions 
in our material relative to the S. parnassum 
standard sequence. Chromosome I is impli- 
cated as the sex chromosome. Two males 
from Georgia (Newsome Gap Road) had 
differentiated centromere regions in chro- 
mosome I; the centromere band of one ho- 
mologue was expanded (standard) more 
than the other (Fig. 5). About 52% of males 
(n = 44) from other sites showed a failure 
to pair on either side of the centromere of 
chromosome I. This failure to pair was 
found at sites with and without the two pu- 
pal forms. One male from Abner Creek (13 
May 1998) was heterozygous for expres- 
sion of the nucleolar organizer. Simulium 
parnassum was otherwise chromosomally 
monomorphic. 


DISCUSSION 


A new pupal form of S. parnassum was 
discovered in which the reticulation of the 
head and thorax, long used as a diagnostic 
character (e.g., Stone and Jamnback 1955), 
was absent. No intermediates between 
smooth and rugose pupae were found; how- 


ee 

symmetrical, 3 = three sharp. 9, Chromosome arm IIIL. Section numbers on bottom indicate S. parnassum 
standard banding sequence; those on top refer to Simulium subgeneric standard sequence of Rothfels et al. 
(1978). Arrows indicate breakpoints of inversions. N.O. = nucleolar organizer, M = marker (divided in two 


pieces). Female from South Carolina, Pickens Co., Oil Camp Creek, 14 May 1998. 


850 


ever, we discovered no additional informa- 
tion, either morphological, chromosomal, 
distributional, or seasonal, to suggest that 
two species are present. The synonymy of 
S. hydationis with S. parnassum (Stone and 
Jamnback 1955) is justified, both morpho- 
logically and cytologically, because mate- 
rial from near the two type localities was 
morphologically and chromosomally ho- 
mogeneous. However, the possibility that 
the two pupal forms of S. parnassum rep- 
resent homosequential species or that hom- 
osequential sibling species exist, as they do 
in other black flies (e.g., Henderson 1986), 
cannot be excluded. 

Relative to the Simulium subgeneric stan- 
dard of Rothfels et al. (1978), we found 
only two inversions that are shared with 
other taxa. The IIS-B inversion of Rothfels 
et al. (1978) is shared with most species in 
subgenus Simulium (Adler et al. 1999), as 
is an inversion in IIIL that has one of its 
breakpoints at the 91/94 junction (Fig. 9) 
(Adler, unpublished). Because IIIL is not 
fully resolved, shared inversions with other 
taxa could be present. The remaining in- 
versions in IL, IIS, and IIL are apparently 
autapotypic for S. parnassum. Chromosom- 
al evidence does not suggest a relation with 
the S. tuberosum species group, which is 
defined by at least eight unique rearrange- 
ments, including four in IIL (Adler and Ku- 
usela 1994), nor does it provide resolution 
of relationship with other species groups. 
Tentative placement of S. parnassum in a 
separate species group, the S. parnassum 
species group, is therefore more appropri- 
ate. 

Simulium parnassum was univoltine in 
South Carolina. In Pennsylvania, it has 
been considered univoltine but with over- 
lapping cohorts (Tessler 1991). Other au- 
thors (e.g., Stone and Snoddy 1969, Cupp 
and Gordon 1983) suggest that S. parnas- 
sum completes more than one generation 
per year, although conclusive evidence sup- 
porting these claims is lacking. 

Based on the results of our study, we 
consider S. parnassum a single species, al- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


beit polymorphic in pupal surface texture. 
We suggest, however, that additional (e.g., 
molecular) evidence be brought to bear to 
corroborate or falsify this hypothesis. 


ACKNOWLEDGMENTS 


We thank D. C. Currie, J. C. Morse, and 
A. G. Wheeler for reviewing the manu- 
script, and C. E. Beard for technical assis- 
tance. This work was supported, in part, by 
grant no. DEB-9629456 from the National 
Science Foundation to PHA. This is Tech- 
nical Contribution No. 4545 of the South 
Carolina Agriculture & Forestry Research 
System. 


LITERATURE CITED 


Addison, E. M. 1980. Transmission of Dirofilaria ursi 
Yamaguti, 1941 (Nematoda: Onchocercidae) of 
black bears (Ursus americanus) by _ blackflies 
(Simuliidae). Canadian Journal of Zoology 58: 
1913-1922. 

Adler, P. H. 1988 [**1987°]. Ecology of black fly sib- 
ling species, pp. 63—76. Jn Kim, K. C. and R. W. 
Merritt, eds., Black Flies: Ecology, Population 
Management, and Annotated World List. Penn- 
sylvania State University, University Park, Penn- 
sylvania, 528 pp. 

Adler, P. H. and K. C. Kim. 1986. The black flies of 
Pennsylvania (Simuliidae, Diptera). Bionomics, 
taxonomy and distribution. Pennsylvania State 
University Agricultural Experimental Station Bul- 
letin 856: 1-88. 

Adler, P. H. and K. Kuusela. 1994. Cytological iden- 
tities of Simulium tuberosum and S. vulgare (Dip- 
tera: Simuliidae), with notes on other Palearctic 
members of the S. tuberosum species-group. En- 
tomologica Scandinavica 25: 439—446. 

Adler, P. H., B. Malmqvist, and Y. Zhang. 1999. Black 
flies (Diptera: Simuliidae) of northern Sweden: 
taxonomy, chromosomes, and bionomics. Ento- 
mologica Scandinavica 29:361—382. 

Crosskey, R. W. and T. M. Howard. 1997. A new tax- 
onomic and geographical inventory of world 
blackflies (Diptera: Simuliidae). The Natural His- 
tory Museum, London, 144 pp. 

Cupp, E. W. and A. E. Gordon. 1983. Notes on the 
systematics, distribution and bionomics of black 
flies (Diptera: Simulidae) in the northeastern 
United States. Search: Agriculture 25: 1-75. 

Davies, D. M. and B. V. Peterson. 1956. Observations 
on the mating, feeding, ovarian development, and 
oviposition of adult black flies (Simuliidae: Dip- 
tera). Canadian Journal of Zoology 34: 615—655. 

Davies, D. M., B. V. Peterson, and D. M. Wood. 1962. 


VOLUME 102, NUMBER 4 


The black flies (Diptera: Simuliidae) of Ontario. 
Part I. Adult identification and distribution with 
descriptions of six new species. Proceedings of the 
Entomological Society of Ontario 92: 70-154. 

Doisey, K. E., R. D. Hall and E J. Fischer. 1986. The 
black flies (Diptera: Simuliidae) of an Ozark 
stream is southern Missouri and associated water 
quality measurements. Journal of the Kansas En- 
tomological Society 59: 133-142. 

Downe, A. E. R. and P. E. Morrison. 1957. Identifi- 
cation of blood meals of blackflies (Diptera: Sim- 
uliidae) attacking farm animals. Mosquito News 
17: 37-40. 

Dyar, H. G. and R. C. Shannon. 1927. The North 
American two-winged flies of the family Simuli- 
idae. Proceedings of the United States National 
Museum 69: 1—54. 

Fuller, H. S. 1940. Black-flies bite woodchuck. Bul- 
letin of the Brooklyn Entomological Society 35: 
155: 

Gibbs, K. E., E C. Brautigam, C. S. Stubbs and L. M. 
Zibilske. 1986. Experimental applications of B.t.i. 
for larval black fly control: persistence and down- 
stream carry, efficacy, impact on non-target in- 
vertebrates and fish feeding. Maine Agricultural 
Experiment Station Technical Bulletin 123: 1—25. 

Henderson, C. A. P. 1986. Homosequential species 2a 
and 2b within the Prosimulium onychodactylum 
complex (Diptera): temporal heterogeneity, link- 
age disequilibrium, and Wahlund effect. Canadian 
Journal of Zoology 64: 859-866. 

Malloch, J. R. 1914. American black flies or buffalo 
gnats. United States Department of Agriculture, 
Bureau of Entomology Technical Series 26: 1—72. 


851 


Rothfels, K. H. 1979. Cytotaxonomy of black flies 
(Simuliidae). Annual Review of Entomology 24: 
507-539. 

Rothfels, K. H. and R. W. Dunbar. 1953. The salivary 
gland chromosomes of the black fly Simulium vit- 
tatum Zett. Canadian Journal of Zoology 31: 226— 
241. 

Rothfels, K. H., R. Feraday, and A. Kaneps. 1978. A 
cytological description of sibling species of Si- 
mulium venustum and S. verecundum with stan- 
dard maps of the subgenus Simulium Davies [sic] 
(Diptera). Canadian Journal of Zoology 56: 1110— 
1128. 

Stone, A. 1964. Guide to the insect of Connecticut. 
Part VI. The Diptera or true flies of Connecticut. 
Ninth Fascicle. Family Simuliidae and Thaumal- 
eidae. State Geological and Natural History Sur- 
vey of Connecticut Bulletin 97: 1—126. 

Stone, A. and H. A. Jamnback. 1955. The black flies 
of New York State (Diptera: Simuliidae). New 
York State Museum Bulletin 349: 1-144. 

Stone, A. and E. L. Snoddy. 1969. The black flies of 
Alabama (Diptera: Simuliidae). Auburn Univer- 
sity. Agricultural Experiment Station Bulletin 390: 
1-93. 

Tessler, S. 1991. Structure of a mountain black fly 
community (Diptera: Simuliidae), Ph.D. Thesis, 
Pennsylvania State University, University Park, 
213 pp. 

Wood, D. M., B. V. Peterson, D. M. Davies, and H. 
Gyorkos. 1963. The black flies (Diptera: Simulii- 
dae) of Ontario. Part II. Larval identification, with 
descriptions and illustrations. Proceedings of the 
Entomological Society of Ontario 93: 99-129. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 852-861 


TYPES OF SAWFLIES DESCRIBED IN THE GENUS PONTANIA A. COSTA 
(HYMENOPTERA: TENTHREDINIDAE) IN THE ILLINOIS 
NATURAL HISTORY SURVEY 


ALEXEY G. ZINOVJEV AND DAVID R. SMITH 


(AGZ) Zoological Institute, Russian Academy of Sciences, St. Petersburg 199034, Rus- 
sia (e-mail: zag@zisp.spb.su); (DRS) Systematic Entomology Laboratory, PSI, Agricul- 
tural Research Service, U.S. Department of Agriculture, % National Museum of Natural 
History, MRC-168, Washington, DC 20560-0168, U.S.A. (e-mail: dsmith @sel.barc.usda. 
gov) 


Abstract.—The types of ten sawfly species described in the genus Pontania by 
MacGillivray and Ross are examined. This study has resulted in the following taxonomic 
changes: Pontania daedala MacGillivray is a new synonym of Pontania proxima (Ser- 
ville); Pontania demissa MacGillivray is a new synonym of Eupontania gracilis (Mar- 
latt); Pontania devincta MacGillivray is a new synonym of Pontania populi (Marlatt); 
Phyllocolpa dedecora (MacGillivray), n. comb.; Phyllocolpa derosa (MacGillivray), n. 
comb.; and Nematus dotata (MacGillivray), n. comb. Notes are given on the types and 


hosts where known. 


Key Words: 


This study of ten sawfly types described 
in the genus Pontania A. Costa deposited 
in the Illinois Natural History Survey, 
Champaign, Illinois (INHS) focuses on 
some that were reared by Yuasa and treated 
by Yuasa (1922) in his work on sawfly lar- 
vae and concurrently or later described by 
MacGillivray (1921, 1923); two species de- 
scribed by Ross (1929) are also included. 
Placement of these species by Smith (1979) 
was based on the generic concepts of that 
time. Study of these specimens is prelimi- 
nary to further work on eastern North 
American gall-forming sawflies and an up- 
dated catalog of Nearctic gall-forming saw- 
flies of the genera Euura Newman, Pontan- 
ia A. Costa, Eupontania Zinovjev, and 
Phyllocolpa Benson. Generic concepts have 
changed since Smith (1979) and the gall- 
forming sawflies of North America are 
poorly known. As a result of extensive 


galls, leaf rolls, Phyllocolpa, Eitelius, Nematus 


work on the Palearctic and world fauna by 
Zinovjev (1993) and Zinovjev and Vikberg 
(in press), the types of Nearctic species 
need to be reexamined and incorporated 
into the current classification. 

We give some notes on each species, il- 
lustrate the types, and attempt to associate 
the MacGillivray species with species treat- 
ed in Yuasa (1922). We propose three new 
synonymies and three new combinations. 
The other four species have been treated 
correctly in previous literature. Species 
headings are in the original combinations. 


SPECIES DESCRIBED BY MACGILLIVRAY 


MacGillivray (1921) described 19 spe- 
cies of Nematinae, including seven species 
of Pontania “‘as a result of an extended se- 
ries of collecting and breeding of sawfly 
larvae by Dr. H. Yuasa at Ithaca, New York, 
and by the Maine Agricultural Experiment 


VOLUME 102, NUMBER 4 


Station at Orono, Maine.” The larvae of all 
these species were supposed to be described 
by Yuasa (1922). We examined part of Yu- 
asa’s collection from Maine (Orono) which 
is deposited in the National Museum of 
Natural History, Smithsonian Institution 
(USNM). It consists of a set of vials with 
larvae or remnants of leaves with galls kept 
in alcohol. 

MacGillivray (1923) described 22 sawfly 
species in a paper on sawflies collected 
from ‘“‘the Katmai Expedition to Alaska.” 
Only one of them was described under the 
genus Pontania and currently (after Benson 
1960) it is treated as a synonym of Phyl- 
locolpa excavata (Marlatt 1896) (= mega- 
cephala Rohwer 1908; destricta Mac- 
Gillivray 1923; apicifrons Malaise 1932; 
carinifrons Benson 1940) (Benson 1960, 
Smith 1979). 


Pontania daedala MacGillivray 1921: 33 
(Ergs2 ie 105418) 


Type locality.—Ithaca, New York. 

Lectotype.—Designated by Frison (1927), 
female, “‘7-6’’; “‘cocoon; Ithaca N. Y. 21 
Aug. 11 [or 17? ]”; “Type of female Pon- 
tania daedala A.D. MacGillivray [red la- 
bell]; “INHS TYPE #1592.” 

Valid name.—Pontania proxima (Ser- 
ville 1823) (= P. daedala MacGillivray), n. 
syn. 

Notes.—The species was reared by Yu- 
asa (1922), and he described the larvae of 
P. proxima (as Pontania hyalina Norton) 
from the material numbered Y-7-1, -7-4-1, 
-8.8, Cu-cu 201, and M-92. The remnants 
of host-plant leaves in the vial (M-92) de- 
posited in the USNM cannot be identified 
to species, but the sawfly is undoubtedly P. 
proxima. We did not find the number ‘‘7- 
6” in Yuasa (1922). 

The female holotype has asymmetric 
mandibles; ovipositor (sawsheath + oblong 
plate) as long as the hindtibia (Fig. 10); 
short cercus, about half the length of the 
sheath; antennal hollow glabrous and shin- 
ing with only few hairs on the lower part 
(Fig. 18), on the upper part the border be- 


853 


tween the inner orbit and antennal hollow 
convex and shining; and supraclypeal area 
glabrous. This combination of characters 
places it in the proxima group. Mac- 
Gillivray mentioned the black color of the 
pronotum which may separate this species 
from P. hyalina Norton (a synonym of P. 
proxima), but in the holotype, the margins 
of the pronotum are pale, exactly as in spec- 
imens of P. proxima from Europe. 


Pontania decrepita MacGillivray 1921: 33 
(Fig. 11) 


Type locality.—Ithaca, New York. 

Holotype female.—‘“Ithaca, N. Y. 21 
July °17; 35-2-5”; “‘Type of female Pon- 
tania decrepita A.D. MacGillivray [red la- 
bel]”; “INHS TYPE #1594.” 

Valid name.—Eitelius gregarius (Marlatt 
1896) (= Pontania decrepita Mac- 
Gillivray). Synonymy by Ross (1951). 

Note.—This species (under the name Mi- 
cronematus gregarius Marlatt) was treated 
by Yuasa (1922) as being collected by him- 
self (““Y’’), but without mentioning its rear- 
ing number, and he did not mention the 
name Pontania decrepita MacGillivray. 
The holotype has claws of the same shape 
as characteristic for Eitelius and also the 
short, broad ovipositor in the holotype (Fig. 
11) is typical for free-feeding Nematinae. 


Pontania dedecora MacGillivray 1921: 32 
ig: 7) 


Type locality—Ithaca, New York. 

Lectotype.—Designated by _ Frison 
(1927), female, designated here; “*185a2, 
May 24, 1919”; *“‘Type of Pontania dedcora 
[sic!] A.D. MacGillivray, female [red la- 
bel]; “INHS TYPE # 1595.”’ 

Valid name.—Phyllocolpa dedecora 
(MacGillivray), n. comb. 

Note.—This species was described from 
two females (““Nos. 185a2, 8.51 (?)-1-1’’). 
Yuasa (1922) mentions none of these num- 
bers. 


854 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Type specimens of Pontania in lateral view. 1, P. daedala. 2, P. demissa. 3, P. devincta. 


VOLUME 102, NUMBER 4 855 


Figs. 4-6. Type specimens of Pontania in lateral view. 4, P. dotata. 5, P. destricta. 6, P. pepii. 


856 


Figs. 7-9. Type specimens of Pontania dorsal view. 


Pontania demissa MacGillivray 1921: 33 
(Figs. 2, 12) 


Type locality.—Ithaca, New York. 

Lectotype.—Designated by Frison (1927), 
female, “‘191-1-1"; “Ithaca N. Y.’’; ‘““Type 
of female Pontania demissa A.D. Mac- 
Gillivray [red label]’’; “INHS TYPE 
FIST.” 

Valid name.—Eupontania gracilis (Mar- 
latt 1896) (= Pontania demissa Mac- 
Gillivray), n. syn. 

Notes.—Larvae and galls were described 
by Yuasa (1922) as P. demissa Mac- 
Gillivray 1921 (Y-191-1-1). We have not 
seen larval material with this number. 

Pontania demissa could be treated either 
as synonym of Eupontania gracilis (Marlatt 
1896) or E. petiolaridis (Rohwer 1917). 
Both these species are associated with dif- 
ferent willow species: Salix sericea Marsh., 
and Salix petiolaris Smith, respectively. 
These sawflies are very similar, and they 
might prove to be conspecific. Both willow 
species occur in the vicinity of Ithaca, and, 
at least in herbarium material studied by 
AGZ, the galls are known also from both 
host plants. However, Salix sericea seem to 
be more common there and galls only on 
this willow were found in the fall of 1997 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


7, P. dedecora. 8, P. derosa. 9, P. mariana. 


around Ithaca by AGZ and T. Carr. The type 
of Pontania demissa fits best the type ma- 
terial of E. gracilis, and we consider them 
as synonymous. 


Pontania derosa MacGillivray 1921: 34 
(Fig. 8, 19) 


Type locality.—Ithaca, New York. 

Holotype.—Female, *‘142-1-1 May 13, 
1919”; “Ithaca, N. Y¥.’’; Dy pexor female 
Pontania derosa A.D. MacGillivray. [red la- 
bell]; “INHS TYPE #1599.” 

Valid name.—Phyllocolpa 
(MacGillivray), n. comb. 

Notes.—The leaf-rolls and larvae were 
described by Yuasa (1922) in his key, but 
the host plant is unknown. The species is 
very similar to Phyllocolpa nigrita Marlatt, 
and these species may be conspecific. 


derosa 


Pontania devincta MacGillivray 1921: 34 
(Figs. 3, 13) 


Type locality—Orono, Maine. 

Lectotype.—Designated by Frison (1927), 
female: ‘““Me. Exp. Sta. Lot. 1694 Sub. 9, 1 
Aug. 13”; “Type of female Pontania de- 
vincta A.D. MacGillivray [red label]”’; 
“INHS TYPE #1601.” 

Valid name.—Pontania populi Marlatt 


VOLUME 102, NUMBER 4 


Figs. 10-13. 
12, P. demissa. 13, P. devincta. 


1896 (= Pontania devincta MacGillivray), 
n. syn. 

Notes.—MacGillivray described this spe- 
cies from two (or more) females, with num- 
bers Subs. 9 and 226. Yuasa (1922) men- 
tioned the number 226, but he did not men- 
tion 9. The galls described by Yuasa, kept 
in the USNM (Maine: Bangor Bay, “*M- 
226’), belong to the gall-making Eupon- 
tania consors (Marlatt 1898). The plant 
species is Salix humilis Marsh. with com- 
paratively large, glabrous leaves (which is 
typical of plants growing in shadowy for- 
ests). We have not seen reared specimens 
with Nr. 226, but according to the galls, the 
paralectotype with this number belongs to 
Eupontania consors. However, the lecto- 
type belongs to the leaf-rolling species 
group of Pontania s. str. and is conspecific 
with Pontania populi, a leaf-rolling species 


857 


Apex of abdomen and sheath in lateral view of Pontania types. 10, P. daedala. 11, P. decrepita. 


on Populus grandidentata Michx. It is con- 
specific also with a specimen identified as 
P. devincta by R. B. Benson (kept in The 
Natural History Museum, London) who 
(1960) incorrectly synonymized Pontania 
devincta under P. excavata Marlatt. 


Pontania dotata MacGillivray 1921: 34 
(Fig. 4) 


Type locality.—Ithaca, New York. 

Lectotype.—Designated by Frison (1927), 
male, “8-48 (?)-1-1”; “Ithaca N.Y. 25 Aug. 
18”; “‘Type of male Pontania dotatus A.D. 
MacGillivray [red label]; ““INHS TYPE 
#1603.” 

Valid name.—Nematus dotatus (Mac- 
Gillivray), n. comb. 

Note.—This species belongs to the genus 
Nematus Panzer (= Pteronidea Rohwer) 
but cannot be placed more precisely at pre- 


858 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 14-17. 14-15, Apex of abdomen and sheath in lateral view of Pontania types. 14, P. destricta. 15, 
P. excavata. 16-17, Head and thorax in lateral view of Pontania types. 16, P. destricta. 17, P. excavata. 


VOLUME 102, NUMBER 4 859 


Figs 18-19. Head in frontal view of Pontania types. 18, P. daedala. 19, P. derosa. 


860 


sent. It seems that this specimen was reared 
by Yuasa, but he did not mention this name 
or the number ‘‘8-48 (?)-1-1” in his keys. 
However, the larvae with a similar number 
(Y-8-48(?)-1) were described by Yuasa as 
Pteronidea sp. 6. 


Pontania destricta MacGillivray 1923: 
168 
(Figs. 5, 14, 16) 


Type locality —Alaska: Katmai 

Holotype.—Female (glued on the card- 
board triangular point): ““Katmai Alaska 
June ’17”’; “Jas S Hine Collector’; Type of 
Pontania destricta A.D. MacGillivray fe- 
male”; “INHS TYPE #1600.” 

Valid name.—Phyllocolpa excavata 
(Marlatt 1896) (= Pontania destricta 
MacGillivray). Synonymy after Benson 
(1960). 

Note.—This species belongs to the Phyl- 
locolpa leucapsis group, characterized by 
glabrous and shining antennal hollows and 
the hairs of the upper posterior part of the 
mesepisternum directed upwards. We are 
not sure that it is conspecific with Phyllo- 
colpa excavata (Marlatt 1896), described 
from ‘California, Colorado” and “*Veta 
Pass, Colo.” The type of P. destricta differs 
from that of Phyllocolpa excavata Marlatt 
1896 (Figs. 15, 17) by a less rounded saw- 
sheath with a distinct emargination beneath 
in lateral view (Figs. 14—15), by the shape 
of the inner orbits in lower part (Figs. 16— 
17), and by the darker coloration (while 
type of P. exavata is paler colored than it 
is typical for this species). 


SPECIES DESCRIBED BY ROSS 


H. H. Ross (1929) described four species 
in his paper revising Marlatt’s “Group I’ 
of Pontania (= Phyllocolpa Benson). Types 
of two of them are deposited in INHS. 


Pontania mariana Ross, 1929: 91 
(i2..9) 
Type locality.—Canada, B.C., Vancou- 
ver. 
Holotype.—Female, reared from _leaf- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


curl on Populus balsamifera; ‘‘Vancouver. 
B. @) Larva coll; x24. 1927 HOH: Ross”: 
‘““emerged VI. 4. 1928”’; [red label] ““HO- 
LOTYPE Pontania mariana Ross female’’; 
“INHS TYPE #1606.” 

Valid name.—Phyllocolpa mariana 
(Ross). 

Host plant.—Populus balsamifera L. 

Additional material examined.—Female 
(paratype), Vancouver. B. C. Larv. Coll. IX. 
4. 1928 H. H. Ross; Reared from Populus 
balsamifera \eaf-curl; emerged VI. 4. 1928; 
[red label] Paratype No 42755; PARATYPE 
Pontania mariana H. H. Ross. Deposited in 
the USNM. 


Pontania pepii Ross, 1929: 95 
(Fig. 6) 


Type locality—NMontana, Florence. 

Holotype.—Female, “‘Mont. Exper. Sta- 
tion Coll. Florence Mont. June 1, 1912”; 
‘“‘“HOLOTYPE Pontania female peppii Ross 
[red label]; ““INHS TYPE #1608.” 

Valid name.—Phyllocolpa pepii (Ross). 


ACKNOWLEDGMENTS 


We thank Kathy Zeiders, Illinois Natural 
History Survey, Champaign, Illinois, for al- 
lowing examination of the MacGillivray 
and Ross types. We extend our appreciation 
to Terry Nuhn for taking the photos and 
Cathy Anderson for arranging the plates. 
Thanks to the following who reviewed the 
manuscript: H. Goulet, Agriculture and 
Agri-Food Canada, Ottawa, and S. Scheffer, 
Systematic Entomology Laboratory, U. S. 
Department of Agriculture, Beltsville, MD. 


LITERATURE CITED 


Benson R. B. 1960. Studies in Pontania (Hym., Ten- 
thredinidae). Bulletin of the British Museum (Nat- 
ural History) Entomology 8(9): 367—384. 

Frison, T. H. 1927. A list of the insect types in the 
collections of the Illinois State Natural History 
Survey and the University of Illinois. State of I- 
linois Department of Registration and Education, 
Division of the Natural History Survey, Bulletin 
16, Article 4, pp. 137-309. 

MacGillivray A. D. 1921. New species of Nemati- 
nae—(Hymenoptera). Journal of the New York 
Entomological Society 29(1): 27-35. 


VOLUME 102, NUMBER 4 


. 1923. Saw-flies of the Katmai expedition to 
Alaska. Journal of the New York Entomological 
Society 31(4): 163-171. 

Marlatt, C. L. 1896. Revision of the Nematinae of 
North America, a subfamily of leaf-feeding Hy- 
menoptera of the family Tenthredinidae. United 
States Department of Agriculture, Technical Se- 
ries No. 3, 133 pp. 

. 1898. Some new nematids. Canadian Ento- 
mologist 30: 302-316. 

Rohwer, S. A. 1917. Pontania petiolaridis, new spe- 
cies, p. 19. In Cosens, A., ed., Reports on Insects 
of the Year, Division No. 3, Toronto District. For- 
ty-Seventh Annual Report of the Entomological 
Society of Ontario, 1916, pp. 18-19. 

Ross H. H. 1929. A study of Marlatt’s group I of the 
genus Pontania with descriptions of four new spe- 
cies (Hymenoptera: Tenthredinidae). Proceedings 
of the Entomological Society of Washington 
31(5): 91-97. 

. 1951. Symphyta, pp. 4-87. In Muesebeck, C. 


861 


FE W., K. V. Krombein, and H. K. Townes, eds., 
Hymenoptera of America North of Mexico, Syn- 
optic Catalog. United States Department of Agri- 
culture, Agriculture Monograph 2, 1420 pp. 

Smith D. R. 1979. Symphyta, pp. 3-137. Jn 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. 

Yuasa H. 1922. A classification of the larvae of the 
Tenthredinoidea. Illinois Biological Monographs 
7(4), 172 pp. 

Zinovjev, A.G. 1993. Subgenera and Palaearctic spe- 
cies groups of the genus Pontania, with notes on 
the taxonomy of some European species of the 
viminalis-group (Hymenoptera: Tenthredinidae). 
Zoosystematica Rossica 2: 145-154. 

Zinovjev A. G. and V. Vikberg. In press. The sawflies 
of the Pontania crassispina-group with a key for 
genera of the subtribe Euurina (Hymenoptera, 
Tenthredinidae, Nematinae). Entomologicae Scan- 
dinavicae. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 862-868 


THE LARVA AND PUPA OF SCHIZONYXHELEA FORATTINI WIRTH AND 
GROGAN (DIPTERA: CERATOPOGONIDAE) WITH A DISCUSSION OF THE 
PHYLOGENETIC RELATIONSHIPS OF THE GENUS 


ART BORKENT 


Research Associate, Royal British Columbia Museum and the American Museum of 
Natural History, 1171 Mallory Road, R1-S20-C43, Enderby, British Columbia, VOE 1V0O, 


Canada (e-mail: aborkent @jetstream.net) 


Abstract.—The larva and pupa of Schizonyxhelea forattinii Wirth and Grogan are de- 
scribed for the first time. Larval, pupal and adult character states suggest that this genus 
is closely related to Stilobezzia Kieffer and may even be most closely related to only 


some species of Stilobezzia. 


Resumen.—Se describen por primera vez larva y pupa de Schizonyxhelea forattinii 
Wirth y Grogan. El estado de los caracteres de larva, pupa y adulto, sugiere que este 
género se halla muy relacionado a Stilobezzia Kieffer, y puede atin, ser mas cercano a 


solo algunas especies de Stilobezzia. 


Key Words: 
Costa Rica, Neotropical 


The immatures of the Ceratopogonidae 
remain as one of the most poorly known 
life stages of all nematoceran families. As 
part of a project to generate descriptions, 
keys and phylogenetic interpretation of 
these stages (Borkent, in preparation), this 
paper provides a description of the larva 
and pupa of Schizonyxhelea forattinii Wirth 
and Grogan and interprets available char- 
acter stages of the genus Schizonyxhelea 
Clastrier from a cladistic perspective. 

Schizonyxhelea includes two Neotropical 
species and both are very small in size, with 
wing lengths of 0.7—0.8 mm. When the ge- 
nus was first proposed by Clastrier (1984), 
he suggested that the single known species, 
S. guyana, was similar to species of Nan- 
nohelea Grogan and Wirth and Bothahelea 
Grogan and Wirth. Wirth and Grogan 
(1988) described a second species, S. for- 
attinii, including the first male of the genus 
and hypothesized that Schizonyxhelea was 


Schizonyxhelea, Stilobezzia, immatures, taxonomy, phylogeny, systematics, 


related to a group of genera with reduced 
features: Baeohelea Wirth and Blanton, 
Baeodasymyia Clastrier and Raccurt, Nan- 
nohelea Grogan and Wirth and Rhynchoh- 
elea Wirth and Blanton. Since then Borkent 
(1992, 1995), Borkent and Craig (1999) and 
Grogan and Borkent (1992) have shown 
that Baeohelea and Baeodasymyia, as sister 
genera, are likely an early lineage within 
the Ceratopogonini and that Nannohelea 
and Rhynchohelea belong to a group which 
also includes the genera Brachypogon Kief- 
fer, Ceratoculicoides Wirth and Ratanawor- 
abhan and Sinhalohelea Grogan and Bor- 
kent. The genus Schizonyxhelea has re- 
mained an enigma. Discovery of the im- 
matures and examination of fresh adult 
material of S. forattinii provided an oppor- 
tunity to further investigate the genus. 


MATERIALS AND METHODS 


Terms for larval and pupal structures fol- 
low Lawson (1951). The single live larva 


VOLUME 102, NUMBER 4 


was discovered by placing a mud sample in 
water and shaking the container gently. The 
swimming larva was then captured with an 
eye-dropper and reared in a petri dish (5 cm 
in diameter) with a small amount of sub- 
strate (mud and detritus) from the original 
habitat. The larva and subsequent pupa 
were kept at ambient temperature (approx- 
imately 25°C) and checked daily. When 
needed, a small amount of water was added 
to the petri dish to keep the sample from 
drying. The larval and pupal exuviae and 
partially emerged adult were preserved in 
70% ethanol and slide-mounted using the 
method described by Borkent and Bissett 
(1990). 

Adults were also captured in the field by 
sweeping vegetation with an aerial net; 
these were examined as cleared specimens 
in glycerine and on slides. Specimens were 
compared to the holotype of S. forattinii. 

The specimens are deposited in the Ca- 
nadian National Collection in Ottawa, On- 
tario, Canada. 


DESCRIPTION OF LARVA AND PUPA 


Fourth instar larva.—Body length un- 
known. Head capsule (Figs. 1A, B) rela- 
tively short (111 pm), blunt (head length/ 
width = 1.28), uniformly medium brown. 
Collar slightly darker than rest of head cap- 
sule, well-developed ventrally with ventro- 
lateral anterior apodeme, separated ventro- 
medially, terminating dorsolaterally. Frontal 
suture extending to near anterior margin of 
labrum. Ventral suture short. Most setae 
simple with the following divided or plu- 
mose: s, v, posterior 0; arrangement as in 
Figs. 1A, B; bases of sensilla more or else 
equal in size; following sensilla not visible: 
j, t, Z, xX, n. Antenna short, further details 
not visible. Eye unknown. Labrum short, 
wide, details not visible. Mandible elon- 
gate, basal portion straight, apical third 
curved, sensilla not visible. Maxilla not 
clearly visible, palpus short. Hypostoma a 
broadly rounded projection. Epipharynx 
with 1 pair of serrate combs, divided me- 
dially (Fig. 1B). Thorax, abdomen with 


863 


more or less uniform light pink pigmenta- 
tion. Caudal segment, anal papillae un- 
known. 

Pupa.—Length 1.29 mm. General color- 
ation uniformly light brown. Body surface 
generally smooth with only a few tubercles 
and spicules associated with setae, a few 
restricted to the medioanterodorsal and me- 
dioanteroventral areas of abdominal seg- 
ments 3—8 and anal segment. Operculum 
(Fig. 1C) with well-developed anteromar- 
ginal seta located on tubercle, with pore at 
tubercle base; dorsolateral margin with row 
of well-developed spines. One well-devel- 
oped anterodorsal seta on moderately elon- 
gate, rounded tubercle (Fig. 1G). One long, 
slender and two shorter dorsolateral setae 
(Fig. 1H). Two short dorsomedial seta, one 
thicker than other (Fig. 1F). Five dorsal 
sensilla, 1 short thick seta, 11, iv short, slen- 
der setae, v, Vi pores; i, ii, 1v on single, 
rounded, short, separate tubercles (Fig. 2A). 
Ventromedian setae not visible (if present, 
they are very minute). Two ventrolateral se- 
tae slender, bases somewhat separate (Fig. 
1E). Respiratory organ (Fig. 1D) somewhat 
curved, smooth; four spiracles situated 
along length, with 5—6 spiracles situated 
apically, opening at surface; tracheal tube 
of more or less equal diameter for length. 
Metathorax not divided medially, with me- 
dial protuberance from scutum protruding 
to near posterior margin. Abdomen with se- 
tae separate from one another (none on 
common tubercle), each bordered by comb- 
like or single elongate spicules. Segment 4 
setal pattern (Fig. 2C) with 2 d.a.s.m., 4 
dp.nisw ls Laisa. 3. pam: 3: yng setae 
d.p.m. i—ili not present or very small; setae 
dipantiv,, Lasimeelipam: 1:, ii, yall thick; 
stout. Anal segment (Fig. 2B) with caudal 
spine moderately elongate, somewhat 
curved apically, directed posterolaterally. 

Bionomics.—The single larva was col- 
lected on December 17, 1993, 2 km north- 
east of Tarcoles, Costa Rica from wet mud 
in a small seep which flowed into the out- 
flow of a larger spring on private property 
directly west of Reserva Carara. The larva 


864 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Mandible 


ZZ, Leppup) Pl) I 


Fig. 1. Structures of Schizonyxhelea forattinii. A, Larval head capsule in dorsal view. B, Larval head capsule 
in ventral view. C, Pupal operculum. D, Pupal respiratory organ. E, Pupal mouthparts in ventral view. E Pupal 
dorsomedial setae. G, Pupal anterodorsal seta. H, Pupal dorsolateral setae. 


swam with the same rapid serpentine mo- two male and 11 female adult S. forattinii. 
tion as do so many Ceratopogoninae. Dur- Each of the sites was at, or very near, a first 
ing eight months in Costa Rica (July, 1993— order seep or spring, or a small stream and 
Feb. 1994) I collected only one larva and specimens were collected from August to 


VOLUME 102, NUMBER 4 865 


7 
| 
IV « 

> 
ii 

. fo) 

B 
vi o A 
Dorsal Ventral 
dasm. ts i 
|.a.S.m. 

|.p.m. hw hw 


Fig. 2. Pupal structures of Schizonyxhelea forattinii. A, Dorsal setae. B, Anal segment in ventral view. C, 
Setae of fourth abdominal segment. 


866 


December (3 sites at 2 km NE Tarcoles 
from Aug. 3 to Dec. 17, 1993; a first order 
spring in Atenas Oct. 1 to Dec. 7, 1993). 
This suggests that immatures of this species 
are restricted to small lotic habitats. 


DISCUSSION 


There are two newly discovered charac- 
ter states which provide cladistic informa- 
tion regarding the phylogenetic position of 
Schizonyxhelea. The first is the presence of 
plumose setae on the head capsule of the 
larva, which merely indicates the unsur- 
prising conclusion that this taxon belongs 
within that group of Ceratopogoninae 
which is the sister group of Ceratopogon 
Meigen and, possibly Baeodasymyia + 
Baeohelea. This character is discussed fur- 
ther by Borkent and Craig (1999). The sec- 
ond character state concerns the nature of 
the male aedeagus. Borkent (1995: 97—98) 
discussed (as his character 39) the uniquely 
divided aedeagus of species of Stilobezzia 
Kieffer. Wirth and Grogan (1988) pointed 
out that the male aedeagus of Schizonyxhe- 
lea was difficult to interpret. However, re- 
examination of fresh material here, espe- 
cially from a posterior aspect, shows that 
the aedeagus of S. forattinii is divided me- 
dially, indicating at least a sister group re- 
lationship to, or within, Stilobezzia. 

Further character states suggest that Schi- 
zonyxhelea is actually closely related to a 
species group within Stilobezzia and they 
are discussed below. Although these fea- 
tures are not yet understood cladistically, 
continuing studies show promise for the in- 
terpretation of most of these (Borkent, in 
preparation). 

—Larval head capsule sensilla q, s, k are 
all close to each other (present in Stilobez- 
zia flavirostris (Winnertz) and S. papillata 
Remm, the only two species of Stilobezzia 
for which this character state has been de- 
scribed). The character state appears to be 
unique within the Ceratopogonini but is 
also present in some Sphaeromiini and Pal- 
pomyiini. 

—The following features are shared with 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


the pupa of Stilobezzia bulla Thomsen: pu- 
pal respiratory organ shape and distribution 
of spiracles virtually identical; presence of 
thick spines on margin of operculum (also 
present in many other genera); same num- 
ber and similar size of thoracic setae; spe- 
cific distribution of sensilla and spines on 
segment 4 especially including the anterior 
lateral sensilla and the presence of comb- 
like spines associated with most sensilla 
very similar; presence of two medial, low 
projections on the dorsum of the anal seg- 
ment; presence of spines on posterior mar- 
gin of apicolateral processes of anal seg- 
ment. 

—AlIl Stilobezzia pupae are missing one of 
the dorsal setae (probably # 3) and this ap- 
pears to be nearly unique within the Cera- 
topogonidae. It is also known in Baeoda- 
symyia; Borkent and Craig (1999) misiden- 
tified sensilla v as iii and vi as v. 

—The male antenna of Schizonyxhelea 
have flagellomeres 5—10 fused. Flagello- 
meres 5—9 are fused in S. bulla and flagello- 
meres 5—10 in S. thomsenae Wirth. The 
males of most species of Stilobezzia have 
separate flagellomeres. 

—A few genera of Ceratopogonini (includ- 
ing Stilobezzia) and most genera of Heter- 
omyiini + Sphaeromiini + Palpomyiini + 
Stenoxenini have a secondary row of pali- 
sade setae (defined in detail by Borkent, in 
press). This is also present in Schizonyxhe- 
lea. 

—Adult eyes of Schizonyxhelea have mi- 
nute interfacetal pubescence. Very small in- 
terfacetal spicules are also present in S. bul- 
la (otherwise rare in Stilobezzia). 

—Wing with radial cells reduced. Although 
Wirth and Grogan (1988) suggested that 
Schizonyxhelea lack radial cells, the male I 
have had a narrow second radial cell. A 
number of Stilobezzia species have a small 
or completely reduced first radial cell. 
However, loss of radial cells is known in a 
number of other genera of Ceratopogonidae 
and the character state cannot be interpreted 
at present. It was likely the reduced radial 
cells which led Wirth and Grogan (1988) to 


VOLUME 102, NUMBER 4 


suggest that Schizonyxhelea was related to 
other genera with reduced wing venation 
but, as noted in the introduction, these taxa 
with reduced character states are not closely 
related to one another. It is apparent that 
several lineages of Ceratopogonidae have 
produced small species which have inde- 
pendently reduced their wing venation and, 
for some, their mouthparts, antennae and 
parts of the male genitalia. 

—Wing with a narrow apical band of ma- 
crotrichia. This character state is shared 
with S. bulla and S. thomsenae and appears 
to be unique with the subgenus Stilobezzia. 
Members of the subgenera Debenhamia 
Wirth and Grogan and Acanthohelea also 
have macrotrichia on the wing membrane 
but this is nearly always more generally dis- 
tributed on the apex of the wing. Stilobezzia 
(Acanthohelea) insolita Das Gupta and 
Wirth from Malaysia also has a narrow 
band of macrotrichia and the female has a 
distinctively (but not unique within the Cer- 
atopogonidae) curved single spermathecae 
very similar to that present in female Schi- 
zonyxhelea. 

—Aedeagus very reduced in size. This 
character state is shared with S. bulla, S. 
thomsenae and a few other Stilobezzia. The 
condition is otherwise rare within the Cer- 
atopogonidae. 

—Male cerci are closely approximated. 
This is present in S. thomsenae, some other 
Stilobezzia and some other genera of Cer- 
atopogonini. 

The striking similarity of the pupae of 
Schizonyxhelea forattinii and Stilobezzia 
bulla and the presence of fusion of male 
flagellomeres 5—9 or 10, the narrow band 
of apical of macrotrichia on the wing and 
the reduced aedeagus shared with S. bulla 
and S. thomsenae probably indicates that 
the two species of Schizonyxhelea are ac- 
tually small Stilobezzia with a somewhat re- 
duced wing venation and with equal claws 
on each leg of the female. A reversion from 
a single claw to double claws is known in 
other genera such as Serromyia Meigen 
(Borkent and Bissett, 1990) and Alluaudo- 


867 


myia Kieffer (personal observation of new 
species) and it would not be surprising to 
find the same within Stilobezzia; indeed, I 
have a small Stilobezzia from Costa Rica 
with equal claws but with typical Stilobez- 
zia-like wing venation (i.e., a small first and 
a long second radial cell is present). 

It is yet possible that the pupal character 
states shared by Schizonyxhelea forattinti 
and Stilobezzia bulla are plesiomorphic 
within Stilobezzia and that Schizonyxhelea 
is therefore the sister group of Stilobezzia. 
Further study is required of character state 
polarities to resolve this question (Borkent, 
in preparation). For the present it is best to 
continue to recognize Schizonyxhelea as a 
valid genus. 


ACKNOWLEDGMENTS 


My wife Annette and brother and sister- 
in-law, Herman and Pieta Borkent, financed 
part of the collecting trip to Costa Rica and 
this is deeply appreciated. Annette also pro- 
vided the resources to work up the results 
of that expedition and I thank her for her 
incredible support. 

El Servicio de Parques Nacionales kindly 
provided permission to collect in the nu- 
merous National Parks and Reserves in 
Costa Rica. I express my appreciation to 
Alvaro Castro, who helped organize our 
lives in Costa Rica and directed me to pris- 
tine habitats on the west coast of Costa 
Rica. Finally, Alvaro Vargas permitted col- 
lecting on his farm, just west of Reserva 
Carara, which included an exceptional vir- 
gin rainforest and some beautiful springs 
from which the larva of Schizonyxhelea for- 
attinii was collected. 

I acknowledge the support of the Global 
Environment Facility of the World Bank 
through the “Biodiversity Resources De- 
velopment Project’? developed by the Na- 
tional Institute of Biodiversity (INBio), the 
National System of Conservation Areas 
(SINAC), Ministry of the Environment and 
Energy (MINAE) in Costa Rica. 

Steve Murphree provided helpful criti- 
cisms as a referee of this paper which was 


868 


much appreciated. My thanks to Nancy E. 
Adams and David G. Furth for the loan of 
the holotype of S. forattinii from the Smith- 
sonian Institution in Washington, D.C. 


LITERATURE CITED 


Borkent, A. 1992. A new key to some genera of Cer- 
atopogonini in the Holarctic (Diptera: Ceratopo- 
gonidae). Entomologica Scandinavica 22: 433- 
436. 

. 1995. Biting Midges in the Cretaceous Amber 

of North America (Diptera: Ceratopogonidae). 

Backhuys Publishers, Leiden, The Netherlands, 

237 pp. 

. In press. Biting Midges (Ceratopogonidae: 
Diptera) from Lower Cretaceous Lebanese Amber 
with a Discussion of the Diversity and Patterns 
Found in Other Ambers. Backhuys Publishers, 
Leiden, The Netherlands. 

Borkent, A. and B. Bissett. 1990. A revision of the 
Holarctic species of Serromyia Meigen (Diptera: 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Ceratopogonidae). Systematic Entomology 15: 
153-217. 

Borkent, A. and D. A. Craig (1999). A revision of the 
Neotropical genus Baeodasymyia Clastrier and 
Raccurt (Diptera: Ceratopogonidae) with a discus- 
sion of their phylogenetic relationships. American 
Museum Novitates 3274: 1—26. 

Clastrier, J. 1984. Schizonyxhelea guyana n.g., n.sp. de 
la Guyane Francaise (Diptera, Ceratopogonidae). 
Revue Francaise d’Entomologie 6: 1—4. 

Grogan, W. L. and A. Borkent. 1992. Sinhalohelea, a 
new genus of predaceous midge from Sri Lanka 
(Diptera: Ceratopogonidae). Proceedings of the 
Entomological Society of Washington 94: 314— 
Bi9% 

Lawson, J. W. H. 1951. The anatomy and morphology 
of the early stages of Culicoides nubeculosus Mei- 
gen (Diptera: Ceratopogonidae = Heleidae). 
Transaction of the Royal Entomological Society 
of London 102: 511—570, pl. 1. 

Wirth, W. W. and W. L. Grogan. 1988. The predaceous 
midges of the world (Diptera: Ceratopogonidae; 
Tribe Ceratopogonini). Flora and Fauna Hand- 
book 4, xv + 160 pp., E.J. Brill. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 869-877 


DESCRIPTION OF THE LAST LARVAL INSTAR AND PUPA OF 
LUCIDOTA ATRA (G. A. OLIVIER 1790) (COLEOPTERA: LAMPYRIDAE), 
WITH A DISCUSSION OF ABDOMINAL SEGMENT HOMOLOGY 
ACROSS LIFE STAGES 


Marc A. BRANHAM AND MIGUEL ARCHANGELSKY 


(MAB) The Ohio State University, Museum of Biological Diversity, 1315 Kinnear Rd., 
Columbus, OH 43212-1192, U.S.A. (e-mail: branham.24 @osu.edu); (MA) Centro Regional 
de Investigaciones Cientificas y Transferencia Tecnologica, CRILAR (CONICET—UNLaR) 
Entre Rios y Mendoza s/n, 5301, Anillaco, La Rioja, Argentina (e-mail: marchangelsky @ 
crilar.com.ar) 


Abstract.—The last larval and pupal stages of the widespread and common North Amer- 
ican firefly, Lucidota atra (G. A. Olivier 1790), are described and illustrated. Last instar 
larvae were collected in rotting logs in the early spring and fed terrestrial snails from the 
same logs until the larvae pupated. The larva of L. atra was misidentifed in the literature 
and has subsequently been misidentified as a species in the genus Photinus. A discussion 
of the homology of abdominal sclerites in larval, pupal, and adult fireflies is provided. 


Key Words: Lampyridae, Lucidota atra, Photinus, larva, pupa, morphology, firefly, 


lightningbug, glowworm, ventrite 


The genus Lucidota, as defined by La- 
porte (1833) and fixed by Motschulsky 
(1853), is restricted to the New World and 
contains some 64 described species. The 
genus ranges from the United States to Ar- 
gentina. Lucidota atra (G. A. Olivier) oc- 
curs from the northeastern United States to 
Central America (McDermott 1966). The 
larva of this species was first described by 
H. E Wickham (1895), but because larvae 
in the tribe Photinini are difficult to distin- 
guish (LaBella and Lloyd 1991), a more de- 
tailed larval description is required. Peter- 
son (1951) apparently incorrectly identified 
the larva on which he based his drawing of 
*“Photinus sp.” in his book “‘Larvae of In- 
sects.’ Upon examining “‘Photinus sp.” in 
Peterson’s larval collection at The Ohio 
State University and comparing it with lar- 
vae reared by one of us (MAB), it was dis- 
covered that Peterson’s larva is actually L. 
atra. 


For this study, last instar larvae were col- 
lected in early spring and kept until eclo- 
sion, thus allowing a positive identification 
from the adult. No larval descriptions exist 
for other species of this genus, most likely 
due to difficulties in rearing firefly larvae 
(Archangelsky and Branham 1998). 


MATERIALS AND METHODS 


Seven last instar larvae were collected in 
a rotting log on April 6, 1993 outside of 
Lawrence, KS, and kept in a glass jar with 
damp wood from the log along with some 
terrestrial snails collected in the same 
wood. Empty snail shells were removed 
from the jar every few days. The wood in- 
side the jar was inspected for moisture con- 
tent periodically. When the wood appeared 
to be drying out, it was moistened with dis- 
tilled water. To further simulate the inside 
of the log, the jar was wrapped with paper 


870 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


to reduce light entering the jar. No special 
requirements were necessary for pupation. 

Three larvae and one pupa were fixed in 
boiling water and transferred to 7O%EtOH. 
In order to study the larval morphology, a 
specimen’s head, mouthparts and antennae 
were dissected, cleared in lactic acid, and 
mounted on microscope slides using Hoy- 
er’s as the mounting medium. The descrip- 
tions and drawings were done using a Wild 
MS dissecting microscope and a Zeiss Ax- 
ioscope 20 compound microscope, both 
with a camera lucida. 


RESULTS 
Lucidota atra (G. A. Olivier) 


Description of last larval instar.— 
Length: 13.0 to 15.0 mm. Body elongate, 
fusiform, slightly flattened dorsoventrally 
(Fig. 1). Whitish ventrally with pink along 
sides of thorax and abdomen. Sclerotized 
regions uniformly light to dark brown and 
granulose. All tergites, except abdominal 
tergites 8 and 9, bearing 3 light colored 
stripes that are more or less parallel to the 
longitudinal axis of body. 

Head capsule: Prognathous, subquadra- 
te, dorsoventraly flattened, and robust (Fig. 
2); retractable within thorax. Labrum and 
clypeus fused. Epicranial suture present as 
well as frontal sutures that extend to bases 
of antennae. One pair of lateral stemmata, 
posterior to base of antennae. Head capsule 
not fused ventrally (Fig. 3). 

Antenna: 3-segmented, partially retract- 
able within membranous base (Fig. 4); orig- 
inating on latero-apical edges of head cap- 
sule. Basal segment widest, attached to 
membranous base, median portion of dorsal 
surface covered with medium length setae 
pointing anteriorly, lateral pointing setae on 
anterior third of segment approximately 2 
to 3 times a long as setae in medial region. Fig. 1. Lucidota atra, fifth instar larva, habitus. 
Second segment shorter than third, narrow- Seale han Senn 
er, evenly covered by long setae, carrying 
a large globular sensorium slightly longer 
than third antennomere. Third segment very 
Short, stout with several short setae, an api- 


VOLUME 102, NUMBER 4 871 


Figs. 2-5. Lucidota atra, head of fifth instar larva. 2, Dorsal view. 3, Ventral view. Scale bar = 1 mm. 4, 
Right antenna, dorsal view. 5, Right mandible, dorsal view. Scale bars = 0.2 mm. 


\ 


SS 


Bil 3 SN 
Zip WN | 4 
ie, ae 14 


Figs. 6-7. 
dorsal view. Scale bar = 0.23 mm. 


cal spine, and a small globular sensorium 
on inner surface just below the antennal 
apex. 

Mandible: Symmetrical, strongly falcate, 
with an inner channel opening subapically 
on outer edge (Fig. 5). Retinaculum present, 
forming 2 inner teeth on the apical third of 
mandible. Basal third of the retinaculum 
covered with a dense brush of setae. Medial 
region of mandible covered by a single row 
of long setae pointing inward toward the 
retinaculum, perpendicular to the inner 
channel of the mandible; 1 long seta par- 
allel to apical point of the mandible, just 
anterior to row of setae located medially. 
One 4-pronged seta or sensory appendage 
on outer margin of mandible, just before 
channel opening; outermost prong of this 
seta longer than the other 3. 

Labium: Closely attached to maxilla, 
formed by a short and strongly sclerotized 
prementum, mentum (distally membranous) 
and submentum (fused to mentum). Pre- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Lucidota atra, fifth instar larva. 6, Labium, dorsal view. Scale bar = 0.15 mm. 7, Left maxilla, 


mentum heart shaped, in both dorsal and 
ventral views with distal apical cleft (Fig. 
6); in dorsal view, bearing 2 basal regions 
of very fine setae with longer setae present 
on the segments of the palp; 2 brushes of 
fine cuticular spines present on each side of 
prementum. Palpus 2-segmented; basal seg- 
ment short, bearing several spines, second 
segment twice as long as first, pointed and 
somewhat forked with a single spine (Fig. 
6). 

Maxilla: Apical region (Fig. 7). Basal re- 
gion (Fig. 3). Long and robust, closely at- 
tached to labium. Cardo (Fig. 3) irregularly 
shaped, bearing no setae. Stipes (Fig. 7) 
very broad, ventral surface covered with se- 
tae and bearing a single long seta; dorsal 
surface bearing 2 long setae. Galea large, 
2-segmented, basal segment very long, 3 
times as long as second segment and lack- 
ing setae; distal segment short, conical and 
bearing several short setae with 1 seta on 
distal apex of segment. Lacinia large, twice 


VOLUME 102, NUMBER 4 


as long as first segment of the galea, inner 
surface covered with a thick brush of cutic- 
ular spines. Palpus 3-segmented, basal seg- 
ment largest, subquadrate, longer than other 
2 segments combined, distal two-thirds 
covered with medium to long setae; second 
segment wider than long and bearing me- 
dium length setae; distal segment subconi- 
cal without setae, bearing a globular sen- 
sorium-type structure. 

Thorax: Prothorax subcircular, wider at 
base, containing retracted head when larva 
is in repose. Meso- and metathorax sub- 
rectangular. Thoracic tergites subdivided by 
sagittal line. Each segment with pleural area 
formed by an upper laterotergite, below it 
an epimeron and episternum separated by 
pleural suture; mesothoracic laterotergite 
subdivided, anterior plate smaller, carrying 
mesothoracic spiracle. Prosternum medium 
sized; meso- and metasterna smaller, nar- 
row, subdivided into an anterior basister- 
num and a posterior sternellum. | pair of 
biforous spiracles present on mesopleuron. 

Legs: 5-segmented, coxae long and cy- 
lindrical, robust; trochanters small, sub- 
triangular in lateral view; femora long and 
cylindrical, widening slightly apically, with 
a single long seta in medial inner portion; 
tibiotarsi as long as femora, tapering to- 
wards distal end; pretarsi strong, simple, 
with a pair of stout setae at base. Double 
row of strong setae on inner margin of ti- 
biotarsi, lacking on inner margin of femora. 

Abdomen: 10-segmented, segments 1 to 
8 similar in shape, tapering toward end; 
each tergite subrectangular, tergites 1 
through 8 divided by a sagittal line and 2 
lighter colored lines parallel to sagittal line; 
lateral portions of tergite 8 lightly colored; 
lateral portions of tergite 9 lightly colored 
and without sagittal line; segment 10 a nar- 
row ring surrounding anal region, carrying 
holdfast organ. Pleural areas well devel- 
oped, segments 1 to 7 subdivided, upper 
plate large, suboval, carrying spiracles, 
lower plate small, narrowly subtriangular; 
pleuron 8 with only 1 suboval plate carry- 
ing a spiracle; pleural areas of segments 9 


873 


and 10 reduced. Abdominal sterna large, 
subquadrate, narrowing towards end of ab- 
domen. Postero-lateral corners of sternite 8 
bearing a twin spotted photic organ. Color 
pattern similar to that of thorax. Biforous 
spiracles present on pleurites 1 to 8. 

Description of pupa.—Female, one day 
old. Slightly curved, ventrally concave; 
young pupa white, older pupa approaching 
charcoal in color. Length: 10.0 to 11.0 mm. 

Head: Completely covered by pronotum 
in dorsal view (Fig. 9), white. Eyes small, 
on sides of head; antennae inserted in front 
of eyes, serrate with 11 obvious segments, 
extending in length to metacoxae; antenna 
and mouthparts white. 

Thorax: Pronotum large, subtriangular, 
slight emargination on either side of ante- 
rior apex, covering head; white or cream. 
Meso- and metanotum shorter than prono- 
tum, subrectangular, carrying wing pads on 
sides; posterior medial portion of mesono- 
tum coming to a point, point lacking on me- 
tanotum. First and second pair of legs fully 
visible in ventral view; third pair of legs 
almost completely covered by wingpads, 
only metatarsus visible. 

Abdomen: Segments wider than long, 
white. Tergite 1 with postero-lateral corners 
pointing perpendicular to sagittal axis of 
pupa; postero-lateral corners of tergites 2 
through 8 coming to a point and directed 
posteriorly. Pleurites fused to the sternites 
(except for pleurite 1 which bears spiracle 
1) thus forming lateral margins of abdom- 
inal “‘ventrites’’ (see Discussion.) First ster- 
nite lacking, first ventrite (sternite 2) par- 
tially visible, remaining ventrites fully vis- 
ible, 7 total in female pupa, (male pupa 
with 8 ventrites total.) Medial-lateral cor- 
ners of ventrite 7 (sternite 8) bearing a twin 
spotted photic organ. 

Spiracles: 9 pairs; First on pleuron of 
mesothorax, remaining 8 on abdominal seg- 
ments | to 8. 


DISCUSSION 


Biology 


The activity period of L. atra adults 
range from early June to July, and the larval 


874 


— 


8 


Figs. 8-9. Lucidota atra, pupa. 8, Ventral view. 9, 


life is suspected to be approximately two 
years long, since both large and small lar- 
vae have been found together during mid- 
summer (Balduf 1935). However, as this 
species has not been successfully reared 
from egg to adult, the two year life cycle 
remains speculative. Because L. atra larvae 
are typically found in rotten logs and 
stumps from the fall through early spring, 
it is assumed that these are the larvae that 
overwinter (Williams 1917; MacDermott 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Dorsal view. Scale bar = 4 mm. 


1964; MAB, personal observation). It 
should be noted, however, that stumps and 
logs are places where many coleopterists 
typically look for beetle larvae. Both larvae 
and pupae produced a glow from a two- 
spotted photic organ when disturbed. The 
two-spotted photic organ is located on the 
eighth sternite of the larvae and seventh 
ventrite (eighth sternite) of the female pupa. 
Therefore, it is expected that male pupa 
would also have a similar organ on its sev- 


VOLUME 102, NUMBER 4 


enth ventrite as the seventh ventrite of adult 
males of this species bare such an organ, 
though the ability to luminesce seems to di- 
minish shortly after eclosion (MAB, per- 
sonal observation). Since no L. atra larvae 
have been found foraging in the open, they 
may be subterranean in habit. Adults usu- 
ally fly during the day, and males follow 
pheromone plumes to the females (Lloyd 
1972). The female is typically up to a third 
larger than the male. 


Traditional Perspective, and Modern View 


Peterson (1951) included a side view 
drawing of a lampyrid larva along with a 
mandible and antennae which was labeled 
“Photinus sp.”’ This same drawing was in- 
cluded by LaBella and Lloyd (1991) and 
has thereafter been used as an example of 
a Photinus larva. Upon comparing our L. 
atra larvae with the actual specimen upon 
which Peterson based his drawing (in the 
Peterson Larval Collection, The Ohio State 
University, Columbus, Ohio), they are iden- 
tical matches. Additionally, Peterson’s de- 
termination label in the vial with this spec- 
imen reads “‘Photinus sp.? .”’ This question 
mark on the determination label, evidently 
put there by Peterson himself, was most 
likely accidentally overlooked, and, thus for 
some 49 years, the specimen has been mis- 
identified as Photinus sp., rather than Lu- 
cidota atra. This mistake is very easy to 
make due to the great morphological simi- 
larity between genera in the tribe Photinini, 
in which both Photinus and Lucidota are 
assigned. The only definitive method to as- 
sociate larvae and adults is to rear the lar- 
vae. 


Abdominal Sclerites in Lampyridae 


Considerable confusion has occurred 
concerning the number of abdominal scler- 
ites in discussions of adult firefly abdomen 
morphology. We believe that this confusion 
has been caused largely by the fact that 
lampyrids posses a varying number of vis- 
ible ventral abdominal sclerites and there is 
a lack of accuracy in defining the terms 


875 


used in descriptions and discussions of the 
abdomen. Without both an understanding of 
the homology among abdominal sclerites 
and the use of accurate terminology, mor- 
phological investigations of lampyrids are 
bound to remain confused. 

The description of both larval and pupal 
states of L. atra is instructive for following 
the reduction and fusion of various abdom- 
inal segments and sclerites from the larval 
stage, where all abdominal sclerites are pre- 
sent and obvious, to the pupal stage, where 
the effect of internalization, reduction and 
fusion can be first detected. In all firefly 
larvae currently known, there are ten ab- 
dominal segments, with the tenth being 
quite small and, therefore, commonly over- 
looked. Each abdominal segment bears a 
tergite, distinct pleurites that bear the spi- 
racles (segments one through eight) and a 
sternite. Identification of abdominal seg- 
ments and sclerites in the larvae is not dif- 
ficult. Reduction of abdominal segments 
and the internalization of sclerites in the 
adults however, can make it difficult to de- 
termine homology among abdominal seg- 
ments. 

In Coleoptera, the adult abdomen is usu- 
ally composed of ten segments in the male 
(with the tenth often being highly reduced 
or fused with the ninth), and nine in the 
female (with the ninth being modified to 
form the genital segment) (Lawrence and 
Britton 1991). As was pointed out by Green 
(1956), the first abdominal segment in adult 
Lampyridae is indicated only by the first 
abdominal tergite, except females of Pho- 
tinus granulatus (Green, p. 597). However, 
investigation concluded that the pleurite 
bearing the first abdominal spiracle is also 
present, though in reduced form, in pupal 
L. atra and the adults of some lampyrids. 
In the adult, the first visible ventral sclerite 
actually is of the second abdominal seg- 
ment, as the ventral portion of the first ab- 
dominal segment is usually so internalized 
and reduced, it is not visible ventrally. This 
condition is termed a “hologastrous type 
abdomen” (Nichols 1989). Additionally, in 


876 


adult lampyrids the abdominal pleurites are 
fused to the sternites, thus forming a con- 
tinuous ventral plate. The median half of 
this ventral plate was the larval sternite and 
the lateral regions on each side were the 
pleurites. Green (1956), therefore suggested 
**... it would be incorrect to refer to the 
ventral segments of the abdomen as ster- 
nites.”’ Lawrence and Britton (1991) use the 
term “‘ventrites’’ to denote sternites that are 
externally visible. 

In light of these two situations: sternite 
one lacking and the fusion of sternites with 
pleurites, while also keeping with Green 
(1956) and Lawrence and Britton (1991), 
we adopt the term “‘ventrites’’ to denote the 
visible ventral sclerites in the L. atra pupa, 
which like other firefly species, has the 
same abdominal morphology as the adult. 
In most firefly species, the female has one 
ventrite fewer than the male, with species 
in the Luciolinae being the exception, the 
male having six and the female having sev- 
en ventrites. McDermott (1964) stated that 
*‘The Lampyridae may be defined as that 
family of the Cantharoidea having usually 
seven visible ventral abdominal segments in 
the male.’’ Apparently, McDermott did not 
count the ninth abdominal segment when 
visible, as a visible ventral segment. This 
may be due to the small size if the ninth 
ventral sclerite in relation to the other ven- 
tral sclerites and the fact that it is usually 
the terminal ventral sclerite. It is our present 
conclusion that the sclerite of the ninth ab- 
dominal segment (ventrite eight) needs to 
be counted as a “‘ventrite’’? when visible. 
Depending upon whether the eighth ventrite 
is concealed under ventrite seven or ex- 
posed, adult males will have either seven or 
eight ventrites (MAB, personal observa- 
tion), with eight ventrites being found in the 
majority of genera family-wide, examined 
by MAB. Therefore, in males of most fire- 
fly species, ventrites one through eight cor- 
respond to abdominal segments two 
through nine. The only known exception is 
for members of the subfamily Luciolinae, 
which bear only six ventrites (McDermott 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


1964; Ballentyne 1987a, b) and the paedo- 
morphic brachypterous male of the Euro- 
pean species Phosphaenus hemiperus La- 
porte (MAB, personal observation). In the 
Luciolinae, the last segment exposed in the 
male is ventrite six (abdominal segment 
seven), with segment eight apparently re- 
duced, or altogether lost, with segment nine 
forming part of the aedeagal sheath which 
is retracted into the abdomen (Ballentyne 
1992). The male of Phosphaenus hemiperus 
more or less retains a larviform type ab- 
domen. Therefore, it is no surprise that Tor- 
re-Bueno’s (Nichols 1989) definition of 
‘“‘sternite = ventrite” is insufficient in con- 
veying the homology of ventral abdominal 
segments in adults of Lampyridae. 

Even though ‘“‘segmental fusion’’ does 
not seem to occur in the firefly abdomen, 
the use of the term ‘“‘ventrite’’ should be 
used with care to avoid confusion of the 
homology of various abdominal segments. 
However, the use of the term “‘ventrite”’ to 
denote only visible abdominal segments in 
the adult, while also keeping in mind (and 
mentioning a point of reference) that ““ven- 
trite one”’’ is actually the ventral sclerite of 
the second abdominal segment (in almost 
all cases), is simply good nomenclature and 
serves to avoid confusion concerning which 
adult abdominal segment is being referred 
to. 


CONCLUSION 


The firefly larva labeled as ‘‘Photinus 
sp.’ by Peterson (1951) is actually the larva 
of Lucidota atra, which is herein rede- 
scribed in greater detail than the original 
description (Wickham 1895) in order to fa- 
cilitate larval identification. Through rear- 
ing this species from larva to adult, it was 
possible to investigate the homology of ab- 
dominal segments and track possible fusion 
or reduction events that lead to a decrease 
in number of visible ventral abdominal 
sclerites in the adult. Fusion of both abdom- 
inal segments and ‘“‘ventrites’’ are not 
known to occur in currently studied lam- 
pyrid taxa. 


VOLUME 102, NUMBER 4 


ACKNOWLEDGMENTS 


We thank Lesley A. Ballentyne for dis- 
cussions concerning Pteroptyx abdominal 
segmentation, and James E. Lloyd, Andrey 
Sharkov, David R. Smith, and John W. 
Wenzel, for their helpful comments on the 
manuscript. 


LITERATURE CITED 


Archangelsky, M. and M. A. Branham. 1998. Descrip- 
tion of the Preimaginal Stages of Pyractomena bo- 
realis (Randall, 1838) (Coleoptera: Lampyridae) 
and Notes on its Biology. Proceedings of the En- 
tomological Society of Washington 100(3): 421— 
430. 

Balduf, W. V. 1935. The Bionomics of Entomophagous 
Coleoptera. John S. Swift, St. Louis, 220 pp. 
Ballentyne, L. A. 1987a. Further Revisional Studies on 
the Firefly Genus Pteroptyx Olivier (Coleoptera: 
Lampyridae:Luciolinae). Transactions of the 

American Entomological Society 113: 117-170. 

. 1987b. Lucioline Morphology, Taxonomy and 

Behaviour: A Reappraisal (Coleoptera: Lampyri- 

dae). Transactions of the American Entomological 

Society 113: 171-188. 

. 1992. Revisional Studies on Flashing Fireflies 
(Coleoptera: Lampyridae), Ph.D. Dissertation, 
University of Queensland, St. Lucia, Brisbane, 
Australia. 

Green, J. W. 1956. Revision of the Nearctic Species of 
Photinus (Lampyridae: Coleoptera). Proceedings 
of the California Academy of Science XXVIII 
(15): 561-613. 


877 


LaBella, D. M. and J. E. Lloyd. 1991. Lampyridae, pp. 
427—428. In Stehr, E W., ed., Immature Insects, 
Vol. 2, Kendall Hunt Publishing Company, Du- 
buque, Iowa, 974 pp. 

Laporte, EL.N. (Comte de Castelnau). 1833. D’une 
Révision du genre Lampyre. Annales de la Société 
Entomologique de France II: 122-153. 

Lawrence, J. F and E. B. Britton. 1991. Coleoptera, 
pp. 543-695. In C.S.I.R.O., ed., The Insects of 
Australia, Second Edition, Vol. 2, Cornell Uni- 
versity Press, Ithaca, New York, 1,137 pp. 

Lloyd, J. E. 1972. Chemical communication in fire- 
flies. Environmental Entomology 1(2): 265-266. 

McDermott, FE A. 1964. The Taxonomy of the Lam- 
pyridae (Coleoptera). Transactions of the Ameri- 
can Entomological Society 90: 1—72. 

. 1966. Lampyridae. Jn Steel, W. O., ed., Co- 
leopterorum Catalogus Supplementa. Pars 9 (edi- 
tio secunda). W. Junk, ‘s-Gravenhage, 149 pp. 

Motschulsky, V. 1853. Etudes Entomologiques I: 24— 
58. 

Nichols, S. W. 1989. The Torre-Bueno Glossary of En- 
tomology. Revised Edition of ‘“‘A Glossary of En- 
tomology” by J.R. de la Torre-Bueno. New York 
Entomological Society, American Museum of 
Natural History, New York, New York, 840 pp. 

Peterson, A. 1951. Larvae of Insects. Part II, Edwards 
Brothers, Inc., 416 pp. 

Wickham, H. F. 1895. On the Larvae of Lucidota, Si- 
noxylon and Spermophagus. Bulletin from the 
Laboratories of Natural History of the State Uni- 
versity of Iowa III, pp. 28-35. 

Williams, E X. 1917. Notes on the Life-History of 
some North American Lampyridae. Journal of the 
New York Entomological Society 25: 11—12. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 878-891 


LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF 
NEASPILOTA PUBESCENS FREIDBERG AND MATHIS (DIPTERA: 
TEPHRITIDAE) ON LESSINGIA FILAGINIFOLIA (HOOKER AND ARNOTT) 
M. A. LANE (ASTERACEAE) IN SOUTHERN CALIFORNIA 


RICHARD D. GOEDEN 


Department of Entomology, University of California, Riverside, CA 92521, U.S.A. 
(e-mail: rgoeden @ucrac1l.ucr.edu) 


Abstract.—Neaspilota pubescens Freidberg and Mathis is a univoltine, monophagous 
fruit fly (Diptera: Tephritidae) developing solely in the flower heads of Lessingia filagin- 
ifolia (Hooker and Arnott) M. A. Lane (Asteraceae) belonging to the subtribe Solidagi- 
ninae of the tribe Astereae in southern California. The egg, second- and third-instar larvae, 
and puparium are described and figured. The anterior thoracic spiracle of the second instar 
has five papillae, reduced to two papillae in the third instar. The second instar has seven 
oral ridges and the third instar eight oral ridges, which, except for the most ventral, eight 
oral ridge in the latter instar, are ventrally toothed. The arrangement of these oral ridges 
in a vertical series lateral to the oral cavity is a distinguishing generic character. The 
larvae feed mainly on the ovules and soft achenes as first and second instars; however, 
as third instars, they may extend their feeding into the receptacle and supplement their 
diet with sap. The nonfeeding prepuparium overwinters in a protective cell that occupies 
much of the excavated flower head and is formed of ovule-, achene-, chaff-, pappus-, and 
corolla-fragments impregnated with excess sap and liquid feces that harden when dry. A 
few prepuparia pupate and emerge from their cells in the late summer and probably 
Overwinter as adults, but most pupariate during the next year in late spring, and emerge 
as adults that aggregate on preblossom host plants to mate and subsequently oviposit. 
Pteromalus sp. (Hymenoptera: Pteromalidae) was reared as a solitary, larval-pupal en- 
doparasitoid from a puparium of N. pubescens. 


Key Words: Insecta, Neaspilota, Lessingia, Asteraceae, nonfrugivorous Tephritidae, bi- 
ology, taxonomy of immature stages, flower-head feeding, monophagy, seed 


predation, parasitoid 


Revision of the genus Neaspilota (Dip- 
tera: Tephritidae) by Freidberg and Mathis 
(1986) facilitated determination of speci- 
mens reared from California Asteraceae 
(Goeden 1989) and stimulated several life- 
history studies, including those on N. viri- 
descens Quisenberry (Goeden and Headrick 
1992), N. wilsoni Blanc and Foote (Goeden 
and Headrick 1999), N. signifera (Coquil- 
lett) (Goeden 2000a), N. aenigma Freidberg 
and Mathis (Goeden 2000b), and N. appen- 
diculata Freidberg and Mathis (Goeden 
2000c). This paper describes some imma- 


ture stages and the life history of a sixth 
species from California, N. pubescens 
Freidberg and Mathis. 


MATERIALS AND METHODS 


The present study was based in large part 
on dissections of flower heads of Lessingia 
filaginifolia (Hooker and Arnott) M. A. 
Lane (Asteraceae) collected during 1990— 
1997 mainly from the following two loca- 
tions in the South and North Sections, re- 
spectively, of the San Bernardino National 


VOLUME 102, NUMBER 4 


Forest: Bautista Canyon at 1,100 m eleva- 
tion, Riverside Co. and North of South Fork 
Campground at 1,550 m, SW San Bernar- 
dino Co. One-liter samples of excised, im- 
mature and mature flower heads containing 
eggs, larvae, and puparia were transported 
in cold-chests in an air-conditioned vehicle 
to the laboratory and stored under refrig- 
eration for subsequent dissection, photog- 
raphy, description, and measurement. Eight 
second- and 12 third-instar larvae and five 
puparia dissected from flower heads were 
preserved in 70% EtOH for scanning elec- 
tron microscopy (SEM). Additional prepu- 
paria and 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 hy- 
drated to distilled water in a decreasing se- 
ries of acidulated EtOH. They were osmi- 
cated for 24 h, dehydrated through an in- 
creasing series of acidulated EtOH and two, 
1-h immersions in hexamethyldisilazane 
(HMDS), mounted on stubs, sputter-coated 
with a gold-palladium alloy, and studied 
and photographed with a Philips XL-30 
scanning electron microscope in the Insti- 
tute of Geophysics and Planetary Physics, 
University of California, Riverside. 

Most adults reared from isolated prepu- 
paria and puparia were individually caged 
in 850-ml, clear-plastic, screened-top cages 
with a cotton wick and basal water reser- 
voir and provisioned with a strip of paper 
toweling impregnated with yeast hydroly- 
zate and sucrose. These cages were used for 
studies of longevity and sexual maturation 
in the insectary of the Department of En- 
tomology, University of California, River- 
side, at 25 + 1°C, and 14/10 (L/D) photo- 
period. Two pairs of virgin males and fe- 
males obtained from emergence cages also 
were held in each of six, separate, clear- 
plastic, petri dishes provisioned with a flat- 
tened, water-moistened pad of absorbant 
cotton spotted with honey (Headrick and 
Goeden 1994) for observations of their 
courtship and copulation behavior. 


879 


Plant names used in this paper follow 
Hickman (1993) and Bremer (1994); te- 
phritid names and adult terminology follow 
Foote et al. (1993). Terminology and tele- 
graphic format used to describe the imma- 
ture stages follow Goeden (2000a, b, c), 
Goeden et al. (1998), Goeden and Headrick 
(1992, 1999), Goeden and Teerink (1997; 
1998; 1999a, b), Teerink and Goeden 
(1999), and our earlier works cited therein. 
Means +SE are used throughout this paper. 
Voucher specimens of N. pubescens im- 
mature stages, adults, and parasitoids reside 
in my research collections. 


RESULTS AND DISCUSSION 
Taxonomy 


Adult.—Neaspilota pubescens was de- 
scribed by Freidberg and Mathis (1986: 55— 
57), who pictured the unpatterned wing, 
along with drawings (p. 56) of the lateral 
aspect of the head; male right foretarsus, 
epandrium, distiphallus, epandrium and cer- 
ci, aculeus and its apex enlarged, and sper- 
matheca. 

Immature stages.—The first-instar larva 
remains undescribed, but the egg, second- 
and third-instar larvae and puparium are de- 
scribed below, as the only stages available 
at this writing. 

Egg: Only three intact eggs were found 
and measured in situ within separate, im- 
mature, preblossom flower heads. These 
eggs were white, opaque, smooth, elongate- 
ellipsoidal, and averaged 0.71 + 0.03 
(range, 0.64—0.74) mm long and 0.18 + 
0.003 (range, 0.17—0.18) mm wide, tapered 
and smoothly rounded at both ends. As no 
eggs were examined by scanning electron 
microscopy, the egg of N. pubescens could 
only be generally compared with the eggs 
of N. viridescens, N. wilsoni, and N. appen- 
diculata, which were described in detail by 
Goeden and Headrick (1992, 1999) and 
Goeden (2000c). 

Second instar: White, elongate-cylindri- 
cal, rounded anteriorly, truncated dorsopos- 
teriorly (Fig. 1A), body segments well-de- 


880 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Det WD Exp 200 jum A... Det WD 


142 64 SE 144 4 


Acc.V Spot MaG 


00k 0 2 g a2 ” - 


Fig. 1. Second instar of Neaspilota pubescens: (A) habitus, anterior to left; (B) gnathocephalon, ventrolateral 
view, |1-minute acanthae, 2-anterior sensory lobe, 3-stomal sense organ, 4-mouthhook, 5-median oral lobe, 6- 
integumental petal, 7-oral ridge; (C) anterior sensory lobe, 1-dorsal sensory organ, 2-terminal sensory organ, 3- 
lateral sensory organ, 4-supralateral sensory organ, 5-pit sensory organ, 6-stomal sense organ; (D) anterior 
thoracic spiracle; (E) caudal segment, l-rima, 2-interspiracular process, 3-intermediate sensory complex; (F) 
intermediate sensory complex, 1-stelex sensillum, 2-medusoid sensillum. 


VOLUME 102, NUMBER 4 


fined, circumscribed anteriorly with few 
minute acanthae (Figs. 1B-1); dorsal sen- 
sory organ well-defined, dome-shaped (Fig. 
1C-1); anterior sensory lobe (Fig. 1B-2, C) 
with terminal sensory organ (Fig. 1C-2), 
lateral sensory organ (Fig. 1C-3), suprala- 
teral sensory organ (Fig. 1C-4), and pit sen- 
sory organ (Fig. 1C-5); stomal sense organ 
(Figs. 1B-3, C-6) ventrolaterad of anterior 
sensory lobe; mouthhook bidentate (Fig. 
1B-4); median oral lobe (Fig. 1B-5), fla- 
belliform, laterally compressed (not 
shown); about six papilliform, integumental 
petals dorsal to each mouthhook (Fig. 1B- 
6); seven oral ridges toothed ventrally, in 
vertical series lateral to oral cavity (Fig. 1B- 
7); prothorax, at least, circumscribed ante- 
riorly by posteriorly-directed, minute acan- 
thae (Fig. 1B-1); anterior thoracic spiracle 
with five, cuboidal papillae (Fig. 1D); lat- 
eral spiracular complexes not seen; caudal 
segment with two stelex sensilla (not 
shown) dorsolaterad and ventrolaterad of 
posterior spiracular plate (Fig. 1E); poste- 
rior spiracular plate bears three ovoid rimae 
(Fig. 1E-1), ca. 0.015 mm long, and four 
interspiracular processes (Fig. 1E-2), each 
with one to four, simple or forked branches, 
longest measuring 0.01 mm; intermediate 
sensory complex (Figs. 1E-3, F) with a ste- 
lex sensillum (Fig. 1F-1) and a medusoid 
sensillum (Fig. 1F-2). 

The habitus of the second instar of N. 
pubescens (Fig. 1A) is more like N. wilsoni 
(Goeden and Headrick 1999), N. signifera 
(Goeden 2000a), N. aenigma (Goeden 
2000b), and N. appendiculata (Goeden 
2000c) than the barrel-shaped second instar 
of N. viridescens (Goeden and Headrick 
1992). The dorsal sensory organ of N. pu- 
bescens is well defined in the second instar 
(Fig. 1C-1), as with N. signifera (Goeden 
2000a) and N. appendiculata (Goeden 
2000c), but is not well defined in N. viri- 
descens (Goeden and Headrick 1992), N. 
wilsoni (Goeden and Headrick 1999), and 
N. aenigma (Goeden 2000b). The integu- 
mental petals of the second instars of all six 
species are papilliform and six in number 


881 


in N. pubescens (Fig. 1B-6), like N. viri- 
descens (Goeden and Headrick 1992), but 
number four in N. signifera (Goeden 
2000a), seven in N. wilsoni (Goeden and 
Headrick 1999) and N. appendiculata (Goe- 
den 2000c) and eight in N. aenigma (Goe- 
den 2000b). In the first instars of all six 
congeners examined to date, though not 
available in the present study, the integu- 
mental petals are broad, flattened, and 
paired (Goeden and Headrick 1992, 1999; 
Goeden 1999a, b). An apparent difference 
in N. pubescens is the five papillae on the 
anterior spiracle of the second instar (Fig. 
1D), compared to eight in N. appendiculata 
(Goeden 2000c), and three to four papillae 
in second instars of N. viridescens (Goeden 
and Headrick 1992), N. wilsoni (Goeden 
and Headrick 1999), N. signifera (Goeden 
2000a), and N. aenigma (Goeden 2000b). 
Finally, the interspiracular processes of N. 
pubescens each bear one to four branches 
like N. aenigma (Fig. 1E-2, Goeden 2000b), 
not two to four branches like WN. signifera 
(Goeden 2000a), nor five to nine branches 
like those of N. viridescens (Goeden and 
Headrick 1992), nor two to six branches 
like those of N. wilsoni (Goeden and Head- 
rick 1999), nor four branches like those of 
N. appendiculata (Goeden 2000c). How- 
ever, it is recognized that most specimens 
of N. pubescens with branch numbers at the 
high end of the range will not differ from 
most other Neaspilota in this character. 
Third instar: Pale yellow, ellipsoidal, 
with posterior spiracular plate dark brown 
to black, tapering anteriorly; posterior spi- 
racular plate on caudal segment flattened 
and upturned dorsally ca. 60° (Fig. 2A), mi- 
nute acanthae circumscribe anterior fifth of 
thoracic and anterior abdominal segments, 
but more common posteriorly (Fig. 2B-1); 
gnathocephalon conical (Fig. 2B); dorsal 
sensory organ an elliptical, flat, poorly de- 
fined pad (Fig. 2C-1) punctured centrally 
and peripherally by pore sensilla (Fig. 2C- 
2); anterior sensory lobe (Fig. 2C) bears ter- 
minal sensory organ (Fig. 2C-3), lateral 
sensory organ (Fig. 2C-4), supralateral sen- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


spot Magn §=Det WD Exp HJ 1mm A 


fal } f 14 


AccV SpotMagn Det WD Exp 
p.00 kV 3.0_174x__ SE_13.4 39 


eas ; 
c= Mar 


Acc.V Spot Magn Det WD. Expl 
0:0 kV 3.0 1309x SE "13.4 47 


Fig. 2. Third instar of Neaspilota pubescens: (A) habitus, anterior to left; (B) gnathocephalon, lateral view, 
1-minute acanthae, 2-oral ridge, 3-stomal sense organ, 4-mouthhook; (C) anterior sensory lobe, 1-dorsal sensory 
organ, 2-pore sensillum, 3-terminal sensory organ, 4-lateral sensory organ, 5-supralateral sensory organ, 6-pit 
sensory organ, 7-integumental petal, 8-stomal sense organ; (D) anterior thoracic spiracle; (E) metathoracic 
(left) and abdominal (right) lateral spiracular complexes, 1-spiracle, 2-verruciform sensilla on metathorax, 


VOLUME 102, NUMBER 4 


sory organ (Fig. 2C-5), and pit sensory or- 
gan (Fig. 2C-6); eight, papilliform (above) 
or spatulate (below), integumental petals in 
two rows above each mouthhook (Fig. 2C- 
7); eight oral ridges (Fig. 2B-2), all but 
most ventral ridge, toothed ventrally and 
lateral to oral cavity; stomal sense organ 
(Figs. 2B-3, C-8) ventrolaterad of anterior 
sensory lobe; mouthhook (Fig. 2B-4) tri- 
dentate (not shown); median oral lobe lat- 
erally flattened, apically pointed (not 
shown); prothorax circumscribed by minute 
acanthae (Fig. 2B-1); verruciform sensilla 
circumscribe prothorax posteriorad of mi- 
nute acanthae (not shown); anterior thoracic 
spiracle on posterior margin of prothorax 
bears two oblong papillae (Fig. 2D); meta- 
thoracic lateral spiracular complex with a 
spiracle (Fig. 2E-1) and three verruciform 
sensillae, one above (not shown) and two 
below the spiracle (Fig. 2E-2); abdominal 
lateral spiracular complex with a spiracle 
(Fig. 2E-3) and two verruciform sensilla 
(Fig. 2E-4) dorsoposteriorad of, and two 
verruciform sensilla ventroposteriorad of, 
the spiracle, these sensilla arranged as two 
vertical pairs; caudal segment circum- 
scribed dorsally by minute acanthae (Fig. 
2F-1); a stelex sensillum dorsolaterad (Fig. 
2F-2), laterad (Fig. 2F-3), and ventrolaterad 
(Fig. 2F-4) of posterior spiracular plate 
(Fig. 2G); each posterior spiracular plate 
(Fig. 2G) bears three ovoid rimae (Fig. 2G- 
1), ca. 0.03 mm in length, and four inter- 
spiracular processes (Fig. 2G-2), each with 
one to three, simple, pointed or forked 
branches, longest branch measuring 0.013 
mm; intermediate sensory complex (Fig. 
2H) with a medusoid sensillum (Fig. 2H-1) 
and a stelex sensillum (Fig. 2H-2). Each 
stelex sensillum surrounding the posterior 
spiracular plate apparently has a single, 


ce 


883 


hemispherical, minute acanthus at its base 
(Fig. 2H-3). 

The habitus of the third instar of N. pu- 
bescens generally is like that reported for 
N. viridescens (Goeden and Headrick 
1992), N. wilsoni (Goeden and Headrick 
1999), N. signifera (Goeden 2000a), and N. 
aenigma (Goeden 2000b), and N. appendi- 
culata (Goeden 2000c). Like N. signifera 
(Goeden 2000a) and N. appendiculata 
(Goeden 2000c), only the anterior part of 
each body segment of N. pubescens is cir- 
cumscribed by minute acanthae, whereas, in 
N. aenigma the anteriors, pleura, and pos- 
teriors of each segment are thus circum- 
scribed (Goeden 2000b); in N. wilsoni, all 
intersegmental areas and all abdominal seg- 
ments except the pleura are circumscribed 
(Goeden and Headrick 1999); and in N. vir- 
idescens, the intersegmental areas are free 
of acanthae (Goeden and Headrick 1992). 
Like N. viridescens (Goeden and Headrick 
1992) and N. wilsoni (Goeden and Headrick 
1999), but not like N. signifera (Goeden 
2000a), N. aenigma (Goeden 2000b) and N. 
appendiculata (Goeden 2000c), the dorsal 
sensory organ is not well defined, and flat- 
tened, and not dome-shaped, in the third in- 
star of N. pubescens (Fig. 2C-1). In the sec- 
ond instar of N. wilsoni (Goeden and Head- 
rick 1999), N. signifera (Goeden 2000a), N. 
appendiculata (Goeden 2000c), and N. pu- 
bescens (Fig. 1C-1) the dorsal sensory or- 
gan is both prominent and dome-shaped, as 
it is in the first instar of all congeners ex- 
cept N. signifera, where it was hidden in 
my specimens and could not be examined 
for comparison (Goeden 2000a), and in the 
present study where first instars were not 
available. 

Additional similarities involved the inte- 
gumental petals in the third instars of all 


3-spiracle, 4-verruciform sensilla on first abdominal segment; (F) anal segment, 1-minute acanthae, 2-dorsolateral 
stelex sensillum, 3-lateral stelex sensillum, 4-ventrolaterad stelex sensillum; (G) posterior spiracular plate, 1- 
rima, 2-interspiracular process, 3-ecdysial scar; (H) intermediate sensory complex, 1-medusoid sensillum, 2- 


stelex sensillum, 3-basal, conical, minute acanthus. 


884 


five congeners examined to date, all of 
which are papilliform and arranged in a 
double row above each mouthhook (Goe- 
den and Headrick 1992, 1999; Goeden 
2000a, b, c). The stomal sense organ of the 
third instar of N. pubescens bears one or 
two verruciform sensilla, two pore sensilla, 
and one or two, cone-shaped or short pa- 
pilliform sensilla (Fig. 2C-8). Thus, it ap- 
pears similar in complexity to the stomal 
sense organ of the second instar (Figs. 1B- 
3, C-6); however, the stomal sense organs 
of the third instars of four congeneric spe- 
cies appear especially well-developed com- 
pared to earlier instars and each bears dif- 
ferent combinations of sensory structures, 
variously described as several cone-shaped 
sensilla in N. viridescens (Goeden and 
Headrick 1992); as papilliform and pit-type 
in N. wilsoni (Goeden and Headrick 1999); 
as verruciform or “‘compound verruciform”’ 
in N. signifera (Goeden 2000a), and ver- 
ruciform and pit-type in N. aenigma (Goe- 
den 2000b). 

The third instars of all six species of 
Neaspilota examined to date have oral ridg- 
es with dentate ventral margins character- 
istically arranged in vertical series ventro- 
laterad of the dorsal sensory organ and lat- 
erad of the oral cavity. Similar oral ridges 
also were described in the second instars of 
N. viridescens (Goeden and Headrick 
1992), N. wilsoni (Goeden and Headrick 
1999), N. signifera (Goeden 2000a), and N. 
appendiculata (Goeden 2000c). The oral 
ridges number eight in N. pubescens (Fig. 
2B-2), seven or eight in the third instar of 
N. aenigma (Goeden 2000b), seven in N. 
appendiculata (Goeden 2000c), but six in 
the second and third instars of the other 
three congeners examined to date. The ap- 
pearance and arrangement of these oral 
ridges appears to be a generic character; 
however, the present study and Goeden 
(2000c) confirm that the oral ridges vary in 
number among third instars of Neaspilota 
species. Also, the most ventral, eighth oral 
ridge of N. pubescens is not ventrally 
toothed (Fig. 2B-2). The third instars of 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Trupanea imperfecta (Coquillett), T. jonesi 
Curran, 7. nigricornis (Coquillett), T. pseu- 
dovicina (Hering), 7. signata Foote, and T. 
wheeleri Curran also bear serrated oral ridg- 
es (Goeden and Teerink 1997b, 1998, 
1999a; Goeden et al. 1998; Knio et al. 
1996; Teerink and Goeden 1999), but these 
oral ridges appear to be fewer in number, 
and are not arranged in a more or less reg- 
ular, vertical row lateral to the mouth hook, 
as in Neaspilota. 

The mouthhooks of the third instars of 
N. appendiculata, N. aenigma, N. signifera, 
N. viridescens, and probably N. pubescens 
(unpublished data), are tridentate (Goeden 
and Headrick 1992; Goeden 2000a, b); 
whereas, those of the third instar of N. wil- 
soni are bidentate (Goeden and Headrick 
1999). Such interspecific differences in 
dentation are supported by our findings that 
the mouthhooks of third-instar Trupanea vi- 
cina are bidentate; whereas, those of 12 
other congeners examined from California 
are tridentate (Goeden and Teerink 2000b 
and citations therein). 

The number and appearance of the stelex 
sensilla surrounding the posterior spiracular 
plate differ among the Neaspilota species 
examined to date. These number only four 
in the first instars of N. wilsoni (Goeden and 
Headrick 1999), N. aenigma (Goeden 
2000b), and N. appendiculata (Goeden 
2000), but, unfortunately, were not ob- 
served with N. signifera (Goeden 2000a) or 
in the present study. This count of stelex 
sensilla remains at four in the second instars 
of N. aenigma (Goeden 2000b), N. appen- 
diculata (Goeden 2000c), and N. pubes- 
cens, increases to six in third instars of N. 
wilsoni (Goeden and Headrick 1999), N. 
aenigma (Goeden 2000b), and N. appendi- 
culata (Goeden 2000c). These stelex sen- 
silla also show inter-instar (intraspecific) 
and interspecific differences in the inci- 
dence and appearance of the minute acan- 
thae that may ring them basally, but this 
was not recognized, studied or recorded by 
my coworkers and me until recently (Goe- 
den 2000b, c; Fig. 2H-2, -3). 


VOLUME 102, NUMBER 4 


pot Magn 
J 3) 2bhy 


wed Det WD Exp -——————__ 50 um Cc 


Acc.V Spots 
RLOLO KS) 475x SE 


Fig. 3. Puparium of Neaspilota pubescens: (A) 
habitus, anterior to left; (B) anterior end, 1-invagina- 
tion scar, 2-anterior thoracic spiracle; (C) caudal seg- 
ment, l-rima, 2-interspiracular process, 3-intermediate 
sensory complex. 


13.2 59 


Puparium: Mostly pale yellow, with pos- 
terior two-three segments grayish to black- 
ened posteriorly, ellipsoidal, and smoothly 
rounded at both ends (Fig. 3A); anterior end 


885 


bears the invagination scar (Fig. 3B-1) and 
anterior thoracic spiracles (Fig. 3B-2); cau- 
dal segment circumscribed by minute acan- 
thae; three stelex sensilla, dorsolaterad, lat- 
eral, and ventrolateral of posterior spiracu- 
lar plates; posterior spiracular plate bears 
three broadly elliptical rimae (Fig. 3C-1), 
and four interspiracular processes, each 
with one to three branches (Fig. 3C-2); in- 
termediate sensory complex (Fig. 3C-3) 
with a medusoid sensillum and a stelex sen- 
sillum. Seven puparia averaged 2.79 + 0.05 
(lange, 255—2:95) mmeainveneth 1.31 
0.06 (range, 1.10—1.60) mm in width. 


DISTRIBUTION AND Hosts 


Freidberg and Mathis (1986) described 
the distribution of N. pubescens as ‘‘South- 
em California south of 35° north latitude 
and west of the Sierra Nevada Mountains.” 
Freidberg and Mathis (1986) and Foote et 
al. (1993) mapped the distribution to in- 
clude only California. 

The only reported and confirmed host 
plant of N. pubescens (reported as appen- 
diculata) is Lessingia (reported as Coreth- 
rogyne) filaginifolia by Goeden (1989), 
which belongs to the subtribe Solidagininae 
of the tribe Astereae in the family Astera- 
ceae (Hickman 1993, Bremer 1994). This 
perennial subshrub has at least two distinct 
varieties and itself is widely distributed in 
coastal scrub, oak woodlands, and grassland 
below 2,600 m throughout California and 
into adjacent southwestern Oregon and 
northern Baja California, Mexico (Hickman 
1993, Shreve and Wiggens 1964), where N. 
pubescens also probably occurs. 

Host records in Freidberg and Mathis 
(1986) present a quandary, as only N. brun- 
neostigmata Doane is reported from Lessin- 
gia (reported as Corethrogyne) filaginifolia, 
including both varieties filaginifolia and 
californica (deCandolle) M. A. Lane (re- 
ported as C. californica). Neither of these 
records obtained from Wasbauer (1972) had 
been checked by Freidberg and Mathis 
(1986). Instead, both varietal records prob- 
ably should refer to N. pubescens, not N. 


886 


brunneostigmata. On the other hand, I have 
only reared N. brunneostigmata from Les- 
singia glandulifera A. Gray and L. lem- 
monii A. Gray, never N. pubescens, which 
is easily distinguished from the former te- 
phritid using the keys in Freidberg and 
Mathis (1986) and Foote et al. (1993). The 
latter host record for N. brunneostigmata 
(as aenigma) was first reported by Goeden 
(1989), and was corrected in Foote et al. 
(1993). The host record for L. glandulifera 
is new. Both represent the only confirmed 
host records for N. brunneostigmata. The 
seven other host records for N. brunneo- 
stigmata listed by Wasbauer (1972) and re- 
peated by Freidberg and Mathis (1986) re- 
main unconfirmed and refer either to un- 
documented records or to probable mis- 
identifications listed in Foote and Blanc 
(1963), which, of course, predated Freid- 
berg and Mathis (1986). In summary, I be- 
lieve that N. pubescens is a true mono- 
phage, and I suspect, the better plant tax- 
onomist in the case whereby C. filaginifolia 
was renamed L. filaginifolia (Hickman 
1993): 


BIOLOGY 


Egg.—lIn each of three, closed, preblos- 
som, immature flower heads of L. filagini- 
folia, a single egg was inserted pedicel-last; 
two of these eggs had their long axes per- 
pendicular to the receptacle, one rested at a 
45° angle to the receptacle. The last egg had 
been inserted through the phyllaries and 
was embedded for half its length in the co- 
rolla of a peripheral floret; whereas, the oth- 
er two eggs were placed between an inner 
phyllary and a peripheral floret. The diam- 
eters of the receptacles of these flower 
heads containing eggs averaged 1.7 + 0.17 
(range, 1.40—1.99) mm. 

Larva.—Upon eclosion, the only two 
first instars found feeding in separate, pre- 
blossom flower heads either tunneled into 
an ovule, or into a corolla before entering 
the ovule to which the corolla was basally 
attached. The receptacles averaged 1.4 mm 
in diameter and an average of 1.5 ovules 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


was damaged in these two flower heads. 
Neither receptacle was abraded or pitted by 
feeding. 

Second instars continued feeding on 
ovules in preblossom flower heads or on 
soft achenes in open, blossom and post 
blossom flower heads. All fed within a se- 
ries of adjacent ovules/soft achenes with 
their bodies more or less perpendicular to 
and their mouthparts directed towards the 
receptacles, but always above the recepta- 
cles. Receptacles of 12 flower heads con- 
taining second instars averaged 1.92 + 0.17 
(range, 1.42—3.42) mm in diameter. These 
flower heads each contained a single larva 
that had destroyed an average of 3.8 + 0.6 
(range, 1—8) ovules/soft achenes. Based on 
23 (range, 17—34) as the average total num- 
ber of ovules. and achenes respectively 
counted in 40 preblossom to postblossom 
flower heads, about 9.5% (range, 2.5—20%) 
of the ovules/soft achenes in the 12 flower 
heads were damaged by second instars. 

Third instars (Fig. 4A) initially continued 
to feed mainly on soft achenes in blossom 
or postblossom flower heads. Twenty-four 
flower heads that averaged 1.95 + 0.14 
(range, 1.10—2.56) mm in diameter each 
contained a single third instar. An average 
of 15 + 2.1 (range, 4-34) of the soft 
achenes therein were damaged, or about 
38% (range, 10—100%) of the average total 
of 40 ovules/soft achenes per flower head. 
These percentages of seed predation per lar- 
va per flower head, like those reported for 
N. aenigma (Goeden 2000b) and N. appen- 
diculata (Goeden 2000c), are on the high 
side among florivorous tephritids studied by 
us to date (Headrick and Goeden 1998). 
This percent seed destruction per flower 
head is exceeded only by gregarious flori- 
vorous species like Trupanea_ conjuncta 
(Adams) (Goeden 1987) and T. pseudovi- 
cina Hering (Goeden and Teerink 1998) or 
by species with large larvae that develop in 
immature or small flower heads like Para- 
cantha cultaris (Coquillett) (Cavender and 
Goeden 1982) and Xenochaeta dichromata 
Snow (Goeden and Teerink 1997a). Also, it 


VOLUME 102, NUMBER 4 


887 


Fig. 4. 
achenes in open flower head, (B) late third instar feeding on soft achenes in open flower head, (C) puparium 
formed in spring by overwintered prepuparium in flower head, (D) mating adults, dorsal view, (E) mating adults, 
ventral view, (F) mating adults lateral view. Lines = 1 mm. 


should be noted that the rates of flower 
head infestation per sample for N. pubes- 
cens was very low, e.g., mean of 4.5% 
(range, 1.5%—10%) for five samples, like 
most Neaspilota in nature that we have 
studied (unpublished data). 

Third instars in flower heads fed with 
their long axes oriented perpendicular to 
and mouthparts directed towards the recep- 
tacles (Fig. 4B). Only five (21%) of the 
third instars in the 24 infested heads scored 
or pitted the receptacles; however, most 
presumably supplemented their diet with 


Life stages of Neaspilota pubescens in Lessingia filaginifolia: (A) early third instar feeding on soft 


sap. Goeden and Headrick (1992, 1999) and 
Goeden (2000c) described and discussed 
this similar type of feeding by N. virides- 
cens, N. wilsoni, and N. appendiculata, re- 
spectively. And, as also reported for all 
three of these congeners that overwinter as 
prepuparia (Goeden and Headrick 1992, 
1999; Goeden 2000c), most third instars be- 
came surrounded for about 90% of their 
lengths by cells, which occupied most of 
the interior of the flower heads and con- 
sisted of ovule-, achene-, chaff-, pappus-, 
and corolla-fragments cemented together by 


888 


liquid feces and sap that hardened when dry 
(Fig. 4C). These protective cells were 
slightly larger than the mature larvae, ex- 
ternally incorporated the outer walls of 
achenes and the few uneaten achenes, and 
were blackened and smooth inside (Fig. 
4C). Upon completing feeding and cell con- 
struction, the larvae oriented with their an- 
terior ends towards the receptacles, retract- 
ed their mouthparts, and formed prepuparia 
(Headrick and Goeden 1998). Most individ- 
uals overwintered in diapause as prepuparia 
(Goeden and Headrick 1992, 1999; Goeden 
2000c; Headrick and Goeden 1998), but a 
few individuals pupariated early and 
emerged in summer (July—August). Prior to 
pupariation the prepuparia reversed their 
orientation within their cells and turned 
180° such that their heads were directed 
away from the receptacles (Fig. 4C). 

Adult.—Adults emerged from overwin- 
tered, mature flower heads, and probably 
are long-lived. Under insectary conditions, 
five unmated males averaged 67 + 23 
(range, 23-152) days, but only two virgin 
females were available for study and aver- 
aged 27 (range, 24—30) days. Such lengthy 
longevities for males compare favorably 
with average adult longevities reported for 
adults of N. viridescens (Goeden and Head- 
rick 1992), N. wilsoni (Goeden and Head- 
rick 1999), N. signifera (Goeden 2000a), N. 
aenigma (Goeden 2000b), and N. appendi- 
culata (Goeden 2000c). 

The premating and mating behaviors of 
N. pubescens were not studied in the field, 
but were observed in petri dish arenas 
found to be so useful with many other non- 
congeneric, nonfrugivorous, tephritid spe- 
cies (Headrick and Goeden 1994). Premat- 
ing behaviors observed with paired N. pu- 
bescens were abdominal pleural distension 
and side-stepping by males while tracking 
females (Headrick and Goeden 1994) and 
rapid wing hamation, sometimes combined 
with lofting about 20° by both sexes (Head- 
rick and Goeden 1994). No trophallaxis or 
nuptial gift presentation was noted as re- 
ported with N. viridescens (Goeden and 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Headrick 1992). Seven matings, six by one 
pair during a 2-week period (Figs. 4D, E, 
F) were observed that began during late af- 
ternoon at dusk and lasted an average of 
1,032 + 73 (range, 850—1,440) min (or 17 
h and 12 min on average). This was three 
to five times longer than the average dura- 
tions of 190 min reported for N. aenigma 
(Goeden 2000b), 235 min reported for UN. 
wilsoni (Goeden and Headrick 1999), 238 
min reported for N. signifera (Goeden 
2000a), 285 min reported for N. appendi- 
culata (Goeden 2000c), and 318 min re- 
ported for N. viridescens (Goeden and 
Headrick 1992). It also was among the lon- 
gest mating durations reported by my col- 
laborators and me (Headrick and Goeden 
1994), exceeded only by a pair of Tephritis 
baccharis (Coquillett) that mated for 2 days 
and nights under similar conditions (Goe- 
den and Headrick 1991). A pair of Dioxyna 
picciola (Bigot) was reported to stay to- 
gether in the field for 28 h, during which 
time they copulated and oviposited three 
times in one day, remained in copula on 
one flower head overnight, and copulated 
and oviposited again three times during the 
following day (Headrick et al. 1996), but 
this involved mate guarding (Headrick and 
Goeden 1994, Headrick et al. 1996), also 
observed with N. signifera (Goeden 2000a). 
As far as could be determined, the extended 
mating by N. pubescens involved continu- 
ous copulation. 

The only precopulary behavior observed 
for the female was an elevation of the ovis- 
cape 20° to 30° coupled with rapid wing 
hamation and lofting, which may have sig- 
naled receptivity (but see postcopulatory 
behavior below). A male was observed to 
initiate mating by rapidly pursuing and 
grabbing the head of the female from the 
front with his forelegs and -tarsi, and wres- 
tling her into submission while he mounted 
her. No copulatory induction behavior was 
noted with N. pubescens and intromission 
was gained rapidly. Most matings were ob- 
served only after copulation had begun, 
suggesting that precopulatory behavior was 


VOLUME 102, NUMBER 4 


perfunctory. The mating position (Figs. 4D, 
E, F) was such that the wings of the male 
were parted about 20°, while the wings of 
the female were parted at about 60°, with 
both pairs of wings centered over their re- 
spective body midlines (Figs. 4D, E). The 
body of the female was parallel to the sub- 
strate, while the body of the male was el- 
evated about 20° anteriorly (Fig. 4F). The 
hind- and midtarsi of the male usually rest- 
ed on the substrate, but sometimes his mid- 
tarsi grasped the base of the oviscape, while 
his foretarsi hooked onto the abdominal 
pleura of the female midway along the ab- 
domen (Figs. 4D, F). The male’s mouth- 
parts were positioned above the second ab- 
dominal tergite of the female (Figs. 4D, FP). 
In this position the male tenaciously held 
onto the female as copulation continued, 
while defeating her increasingly vigorous 
efforts to dislodge him. These dislodgment 
efforts by the female consisted of walking 
rapidly, then stopping to strongly arch her 
dosum at the juncture of her thorax and ab- 
domen while rapidly hamating and lofting 
her wings (Fig. 5A), or bending her ovi- 
scape upward, while kicking at his head, 
ventrum, and mid- and hind legs with her 
hind legs and tarsi. In between these bouts 
of female “restlessness,” the pair largely 
remained quiescent with only their mouth- 
parts pumping, or sometimes both forming 
regurgitation droplets (Headrick and Goe- 
den 1994), or while the female groomed her 
head and fore legs. Besides tightly clinging 
to the female, the male sometimes reposi- 
tioned his foretarsi so as to gain better pur- 
chase or rub his midtarsi alternately along 
her oviscape to induce passivity as a vari- 
ation on copulatory induction behavior 
(Headrick and Goeden 1994). These bursts 
of activity by the female always preceded 
disengagement, but only infrequently re- 
sulted in disengagement. What action(s) fi- 
nally triggered or caused the pair to sepa- 
rate, remain unanswered. 

Two mating terminations and disengage- 
ments were observed, which involved each 
male turning in place and walking away 


Figs 5: 
mating by Neaspilota pubescens: (A) female arching 
dorsum and kicking at male with hind legs and tarsi, 
(B) male turning and stepping off female, (C) male 
pulling genitalia from female as he walks away. Lines 
= 1 mm. 


Disengagement sequence at termination of 


from the female while pulling his genitalia 
from within her, a process that lasted less 
than 10 s in both cases (Figs. 5A, B. C). 
Postcopulatory behavior by N. pubescens 
mainly consisted of storing of the genitalia 
by males and cleaning and grooming by 
both sexes (Headrick and Goeden 1994); al- 


890 


though, instances of a female strongly ele- 
vating her oviscape was seen, as she ran 
away from a pursuing male to avoid recou- 
pling following disengagement. As noted 
above, this same behavior often preceded 
mating. 

Seasonal history.—The life cycle of N. 
pubescens in southern California follows an 
aggregative pattern (Headrick and Goeden 
1994, 1998) in which the prepuparium is 
the principal overwintering stage. Come 
late spring (May), overwintered prepuparia 
reverse their orientation in their cells in 
flower heads on shoots of dead host plants 
and pupariate. Adults emerge during late 
May and early June and aggregate on pre- 
blossom shoots of L. filaginifolia to mate. 
Females oviposit in the small, newly- 
formed, closed, preblossom flower heads in 
July and larvae feed until fully grown, then 
enter diapause in early fall (September—Oc- 
tober). There is a single generation per year 
on their sole host plant, although as men- 
tioned above, a few adults emerge in late 
August-September, perhaps to produce a 
partial generation on late-flowering plants, 
or to overwinter as long-lived adults. 

Natural enemies.—A single female of 
Pteromalus sp. (Hymenoptera: Pteromali- 
dae) was reared from a puparium of N. pu- 
bescens as a solitary, larval-pupal endopar- 
asitoid. 


ACKNOWLEDGMENTS 


I thank Andrew C. Sanders, Curator of 
the Herbarium, Department of Botany and 
Plant Sciences, University of California, 
Riverside, for identifications of plants men- 
tioned in this paper. Krassimer Bozhilov in 
the Institute of Geophysics and Planetary 
Physics, University of California, River- 
side, greatly facilitated my scanning elec- 
tron microscopy. The parasitoid was iden- 
tified by Harry E. Andersen, Huntington 
Beach, California. I also am grateful to Jeff 
Teerink and Kristine Gilbert for technical 
assistance and to Louie Blanc, David Head- 
rick, and Jeff Teerink for their helpful com- 
ments on earlier drafts of this paper. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


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. and EF L. Blanc. 1963. The fruit flies or 
Tephritidae of California. California Insect Survey 
Bulletin 7: 1-117. 

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. 

Freidberg, A. and W. N. Mathis. 1986. Studies of Ter- 
elliinae (Diptera: Tephritidae): A revision of the 
genus Neaspilota Osten Sacken. Smithsonian 
Contributions to Zoology 439: 1—75. 

Goeden, R. D. 1987. Life history of Trupanea con- 
jJuncta (Adams) on Trixis californica Kellogg in 
southern California (Diptera: Tephritidae). The 
Pan-Pacific Entomologist 63: 284-291. 

. 1989. Host plants of Neaspilota in California 

(Diptera: Tephritidae). Proceedings of the Ento- 

mological Society of Washington 91: 164—168. 

. 2000a. Life history and description of im- 

mature stages of Neaspilota signifera (Coquillett) 

(Diptera: Tephritidae) on Hemizonia pungens 

(Hooker and Arnott) Torrey and A. Gray (Aster- 

aceae) in southern California. Proceedings of the 

Entomological Society of Washinton 102: 69-81. 

. 2000b. Life history and description of im- 

mature stages of Neaspilota aenigma Freidberg 

and Mathis (Diptera: Tephritidae) on Erigeron 
divergens Torrey and Gray (Asteraceae) in south- 
ern California. Proceedings of the Entomological 

Society of Washington 102: 384-397. 

. 2000c. Life history and description of im- 
mature stages of Neaspilota appendiculata Freid- 
berg and Mathis (Diptera: Tephritidae) on Ma- 
chaeranthera canescens (Pursh) A. Gray (Aster- 
aceae) in southern California. Proceedings of the 
Entomological Society of Washington 102: 519— 
DBZ: 

Goeden, R. D. and D. H. Headrick. 1991. Life history 
and descriptions of immature stages of Tephritis 
baccharis (Coquillett) on Baccharis_salicifolia 
(Ruiz & Pavon) Persoon in southern California 
(Diptera: Tephritidae). The Pan-Pacific Entomol- 
Ogist 67: 86-98. 

. 1992. Life history and descriptions of imma- 

ture stages of Neaspilota viridescens Quisenberry 

(Diptera: Tephritidae) on native Asteraceae in 

southern California. Proceedings of the Entomo- 

logical Society of Washington 94: 59-77. 

. 1999. Life history and description of imma- 

ture stages of Neaspilota wilsoni Blanc and Foote 


VOLUME 102, NUMBER 4 


(Diptera: Tephritidae) on Hazardia squarrosa 
(Hooker and Arnott) E. Greene (Asteraceae). Pro- 
ceedings of the Entomological Society of Wash- 
ington 101: 897-909. 

Goeden, R. D. and J. A. Teerink 1997a. Life history 
and description of immature stages of Xenochaeta 
dichromata Snow (Diptera: Tephritidae) on Hier- 
acium albiflorum Hooker in central and southern 
California. Proceedings of the Entomological So- 
ciety of Washington 99: 597-607. 

- 1997b. Life history and description of im- 

mature stages of Trupanea signata Foote (Diptera: 

Tephritidae) on Granphalium luteo-album L. in 

southern California. Proceedings of the Entomo- 

logical Society of Washington 99: 748-755. 

. 1998. Life history and description of imma- 

ture stages of Trupanea pseudovicinia Hering 

(Diptera: Tephritidae) on Porophyllum gracile 

Bentham (Asteraceae) in southern California. Pro- 

ceedings of the Entomological Society of Wash- 

ington 100: 361-372. 

. 1999a. Life history and description of im- 

mature stages of Trupanea wheeleri Curran (Dip- 

tera: Tephritidae) on Asteraceae in southern Cal- 
ifornia. Proceedings of the Entomological Society 

of Washington 101: 414—427. 

. 1999b. Life history and description of im- 
mature stages of Trupanea vicina (Wulp) (Diptera: 
Tephritidae) on wild and cultivated Asteraceae in 
southern California. Proceedings of the Entomo- 
logical Society of Washington 101: 742-755. 

Goeden, R. D., J. A. Teerink, and D. H. Headrick. 
1998. Life history and description of immature 
Stages of Trupanea jonesi Curran (Diptera: Te- 
phritidae) on native Asteraceae in southern Cali- 


891 


fornia. Proceedings of the Entomological Society 
of Washington 100: 126-140. 

Headrick, D. H. and R. D. Goeden. 1994. Reproduc- 
tive behavior of California fruit flies and the clas- 
sification and evolution of Tephritidae (Diptera) 
mating systems. Studia Dipterologica 1(2): 194— 
Wy. 

. 1998. The biology of nonfrugivous tephritid 
fruit flies. Annual Review of Entomology 43: 
217-241. 

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 Coreopis spp. (Asteraceae) in south- 
ern California. Proceedings of the Entomological 
Society of Washington 98: 332-349. 

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. 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. 

Shreve, F and I. L. Wiggens. 1964. Vegetation and 
Flora of the Sonoran Desert, Vol. 2. Stanford Uni- 
versity Press, Stanford, California. 

Teerink, J. A. and R. D. Goeden. 1999. Description of 
the immature stages of Trupanea imperfecta (Co- 
quillett). Proceedings of the Entomological Soci- 
ety of Washington 101: 75-85. 

Wasbauer, M. S. 1972. An annotated host catalog of 
the fruit flies of America north of Mexico (Dip- 
tera: Tephritidae). Bureau of Entomology, Cali- 
fornia Department of Agriculture Occasional Pa- 
per 19: 1-172. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 892-900 


A TAXONOMIC REVIEW OF THE GENUS DELPHINIOBIUM MORDVILKO 
(HOMOPTERA: APHIDIDAE) IN CHINA 


GE XIA QIAO AND GUANG XUE ZHANG 


Institute of Zoology, The Chinese Academy of Sciences, 19 Zhongguancun Lu, Haidian, 
Beijing 100080, People’s Republic of China (e-mail: leifm@panda.ioz.ac.cn) 


Abstract.—The aphid genus Delphiniobium Mordvilko from China is reviewed. There 
are four species in China, Delphiniobium gyamdaense Zhang, D. yezoense Miyazaki, D. 
violisuctum Qiao and Zhang, n. sp., and D. aconitifoliae Zhang and Qiao, n. sp. A 
key to species from China, morphological descriptions, distributions, host plants, and 
diagnosis of new species are provided. In addition, the apterous viviparous female of D. 
gyamdaense Zhang is described for the first time. 


Key Words: 


Mordvilko (1914) erected the genus Del- 
phiniobium for the species Myzus junack- 
ianum Karsch 1887. Hille Ris Lambers 
(1947) studied the genus Delphiniobium 
from Europe. At present, this genus is rep- 
resented by eight species and one subspe- 
cies, D. bogdouli Szelegiewicz 1969, D. 
canadense (Robinson 1968), D. carpaticae 
Mamontova 1966, D. hanla Paik 1971, D. 
Junackianum (Karsch 1887), D. junackinum 
sylvanae (Knechtel and Manolache 1944), 
D. lycoctoni B6rner 1950, D. yezoense Mi- 
yazaki 1971 and D. gyamdaense Zhang 
1981 (Remaudiére and Remaudiére 1997). 
In the course of the present study, two new 
species, D. aconitifoliae Zhang and Qiao 
and D. violisuctum Qiao and Zhang are de- 
scribed from China, plus two other species, 
D. gyamdaense Zhang and D. yezoense Mi- 
yazaki. A key to species in China is pro- 
vided. In addition, the apterous viviparous 
female of D. gyamdaense Zhang 1981 is 
described for the first time. 


MATERIALS AND METHODS 


All specimens studied including types are 
housed in the Zoological Museum, Institute 


Homoptera, Aphididae, Delphiniobium, new species, China 


of Zoology, the Chinese Academy of Sci- 
ences, Beijing. 

Aphids were mounted on microscope 
slides in Arabic gum, and observed under 
phase contrast microscopy. Terminology 
follows Miyazaki (1971). Drawings were 
made by the first author, Dr. X. L. Chen, 
and Mr. T. S. Zhong using microscopy. 
Measurements are in millimeters. 


Delphiniobium Mordvilko 1914 


Delphiniobium Mordvilko 1914: 65. 

Type-species: Myzus junackianum Karsch 
1887 (= Rhopalosiphum aconiti van der 
Goot 1912). By monotypy. 


Generic diagnosis.—Body elliptical. Me- 
dial frontal tubercle undeveloped or devel- 
oped, lateral frontal tubercles with diverg- 
ing inner sides present. Thoracic stigmal 
pores much longer than abdominal stigmal 
pores. Spiracular sclerites of thorax strong- 
ly produced, with opening very large and 
round. Mesosternal furca with short or long 
stem. Antennal segment III with large and 
small round secondary rhinaria. Ultimate 
rostral segment stout. First tarsal segment 
chaetotaxy: 3, 3, 3. Siphunculi slender, 


VOLUME 102, NUMBER 4 


slightly or distinctly swollen, with distinct 
reticulation consisting of several rows of 
hexagonal cells near apex, paler than cauda 
basally. Cauda dark, long tapering. Mainly 
on Aconitum and Delphinium. 

In general aspects this genus resembles 
Megoura Buckton, from which it can be 
easily separated by the reticulated apices of 
the siphunculi. On the other hand many 
Macrosiphum-like species in America have 
swollen siphunculi with a reticulated area, 
but in none of them a black, sclerotic cauda 
occurs. Other genera with swollen siphun- 
culi have either no reticulated area (Ampho- 
rophora Buckton), or (Rhopalosiphoninus 
Baker) often no rhinaria on the antennal 
segment III in apterous viviparous females. 

Distribution.—Europe (Romania, Eng- 
land, Netherlands, Germany, Russia), North 
America (Canada), Asia (China, Japan, Ko- 
rea, Mongolia). 


KEY TO APTEROUS VIVIPAROUS FEMALES OF 
SPECIES FROM CHINA 


1. Medial frontal tubercle not developed, antennal 
tubercles small (Fig. 22); abdominal tergite I 
with 14 hairs; ultimate rostral segment with 2 
pairs accessory hairs; China: Shanxi (Yang- 
cheng County) Delphiniobium violisuctum 
Qiao and Zhang, n. sp. 
— Medial frontal tubercle developed, antennal tu- 
bercles large (Figs. 1, 15); abdominal tergite I 
with at most 10 hairs; ultimate rostral segment 
with 3—5 pairs accessory hairs ........... 2 
. Siphunculi slightly or distinctly swollen in 
middle; ultimate rostral segment at most 1.30 
times as long as second hind tarsal segment; 
antennal segment III with either at most 25 sec- 
ondary rhinaria or at least 44............. 3 
— Sipunuculi not swollen; ultimate rostral seg- 
ment 1.35 times as long as second hind tarsal 
segment; antennal segment III with 21—39 sec- 
ondary rhinaria; China: Beijing (Sanpu Dis- 
trict, Xiaolongmen District) 
Delphiniobium aconitifoliae 
Zhang and Qiao, n. sp. 
3. Antennal segment HI with 44—57 secondary 
rhinaria, on basal %4 of segment; siphunculi dis- 
tinctly swollen (Fig. 20); China: Qinghai (Hu- 
zhu County), Xizang (Gyamda County) 
Bre ori Bers Delphiniobium gyamdaense Zhang 
— Antennal segment III with 10—25 secondary 
rhinaria, on basal half of segment; siphunculus 
slightly swollen; China: Sichuan (An County), 


N 


893 


Hebei (Kuancheng County, Wulinshan Moun- 
tain, Xiaowutai Mountain); Japan ....... 
Ae TE ee Meta Delphiniobium yezoense Miyazaki 


KEY TO ALATE VIVIPAROUS FEMALES OF 
SPECIES FROM CHINA 


1. Siphunculi distinctly swollen; antennal seg- 


ment III longer than segmentsIV+V..... 2 
— Siphunculi indistinctly swollen; antennal seg- 
ment III shorter than segments IV + V .... 3 


N 


. Antennal segment III with 77-81 secondary 
rhinaria; yellow in life; siphunculus 2.00 times 
as long as cauda; China: Qinghai (Huzhu 
County), Xizang (Gyamda County) ..... 
Delphiniobium gyamdaense Zhang 
— Antennal segment HI with 40—65 secondary 
rhinaria; bluish green in life; siphunculus 1.60— 
1.70 times as long as cauda; China: Hebei (Ku- 
ancheng County, Wulinshan Mountain, Xia- 
owutai Mountain), Sichuan (An County); Japan 
Delphiniobium yezoense Miyazaki 
3. Median frontal tubercle indistinct; ultimate ros- 
tral segment 1.06 times as long as second hind 
tarsal segment; siphunculi 1.74 times as long 
as cauda; China: Shanxi (Yangcheng County) 
Delphiniobium violisuctum 
Qiao and Zhang, n. sp. 
— Median frontal tubercle distinct; ultimate ros- 
tral segment 1.40 times as long as second hind 
tarsal segment; siphunculi 2.00 times as long 
as cauda; China: Beijing (Sanpu District, Xiao- 
longmen District) . . 


Delphiniobium aconitifoliae 
Zhang and Qiao, n. sp. 


Delphiniobium aconitifoliae Zhang and 
Qiao, new species 
(Figs. 1-14) 


Description.—Measurements: Apterous 
viviparous female: Body 3.56 in length, 
1.78 in width. Antenna 3.72, length of seg- 
ments I-VI 0.15, 0.09, 1.20, 0.59, 0.50, 
0.12+1.08, respectively. Ultimate rostral 
segment 0.19 in length, 0.09 in basal width. 
Hind femur 1.41, hind tibia 2.53, second 
hind tarsal segment 0.16. Siphunculus 0.72 
in length. Cauda 0.50 in length. Alate vi- 
viparous females: Body 3.20 in length, 1.18 
in width. Antenna 3.93, length of segments 
EVE (O41 45. OOH WIAD) 0262; .0:58; 
0.12+1.23, respectively. Ultimate rostral 
segment 0.21 in length, 0.11 in basal width. 
Hind femur 1.38, hind tibia 2.51, second 
hind tarsal segment 0.15. Siphunculus 0.72 
in length. Cauda 0.37 in length. Alate male: 


894 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-14. Delphiniobium aconitifoliae. 1-9, Apterous viviparous female. 1, Dorsal view of head. 2, An- 
tennal segments I-III. 3, Antennal segment VI. 4, Ultimate rostral segment. 5, Dorsal view of abdominal seg- 
ments V—VIII. 6, Dorsal hair of body. 7, Ventral hair. 8, Siphunculus. 9, Cauda. 10—11, Alate viviparous female. 
10, Antennal segment III. 11, Fore wing. 12-13, Alate male. 12, Antennal segments III-V. 13, Clasper. 14, 
Oviparous female, hind tibia. 


Body 2.64 in length, 1.13 in width. Ovip- 
arous female: Body 3.61 in length, 1.83 in 
width. 

Apterous viviparous female: Body oval, 
red and yellow in life. Mounted specimens: 
Head grey, thorax and abdomen pale, with- 
out patches. Antenna black except base of 
antennal segment III pale; rostral segments 
III-V, distal half of femora, tibiae and tarsi 
black, siphunculi pale at base, brown at 
middle, black distally, cauda and anal plate 
black, genital plate pale. Abdominal seg- 
ments VII and VIII with slight imbrications. 
Spiracles large, closed, spiracular plates 
slightly brown. Mesosternal furca with 
short stem, pale. Dorsal hairs of body thick 


and short, stout at apex. Ventral hairs sim- 
ilar to but longer than dorsal hairs. Head 
with 1 pair cephalic hairs, 1 pair antennal 
tubercular hairs, 4—5 pairs dorsal hairs. 
Pronotum with 1 pair spinal, 1 pair pleural 
and 1 pair marginal hairs. Abdominal ter- 
gites I-VI each with 4—6 spino-pleural 
hairs, tergite VII with 2 or 3; tergites II- 
VII each with 2-3 pairs marginal hairs, ter- 
gite I with | pair marginal hairs, tergite VIII 
with 2 pairs hairs. Length of cephalic hairs 
0.04, length of marginal hairs on abdominal 
tergite I 0.03, and length of dorsal hairs on 
tergite VIII 0.05, 0.52 times, 0.39 times and 
0.75 times as long as widest diameter of 
antennal segment III, respectively. Median 


VOLUME 102, NUMBER 4 


frontal tubercle developed, antennal tuber- 
cles developed and diverging, higher than 
median front. Antenna 6-segmented, basal 
part of segment III and segments IV—VI 
with distinct imbrications, length 1.10 times 
as long as body; length in proportion to seg- 
ments I-VI: 13, 7, 100, 49, 41, 10+90, re- 
spectively. Antennal hairs similar to dorsal 
hairs, segments I-VI each with 6 or 7, 4, 
25-31, 7-13, 7-10, 3 or 4 +3-8 hairs, re- 
spectively, apex of processus terminalis 
with 3 hairs. Length of hairs on antennal 
segment III 0.04, 0.57 times as long as wid- 
est diameter of segment. Antennal segment 
III with 21—39 large and small round sec- 
ondary rhinaria (Fig. 2) on basal % of seg- 
ment. Rostrum thick and large, reaching 
midcoxae, clypeus with 2 pairs hairs ante- 
riorly; ultimate rostral segment 2.00 times 
as long as its basal width, 1.20 times as 
long as second hind tarsal segment, with 7 
pairs hairs, 3—4 pairs accessory hairs 
among them. Hind femur 1.20 times as long 
as antennal segment III. Hind tibia 0.71 
times as long as body. Length of hairs on 
hind tibia 0.06, 0.93 times as long as middle 
tibia width of segment. First tarsal segment 
chaetotaxy: 3, 3, 3. Siphunculi long, taper- 
ing, slightly constricted at distal %4, with 
12-14 rows of reticulations at constricted 
part; 0.20 times as long as body, 1.40 times 
as long as cauda; with flange. Cauda taper- 
ing, rough, with spinulose imbrications; 
with 6-8 thick and long hairs. Anal plate 
circular at apex, with 11—19 long and short 
hairs. Genital plate semicircular, with 14— 
16 hairs. 

Alate viviparous female: Mounted spec- 
imens: Head and thorax slightly brown, 
head surrounding eyes black, abdomen 
pale, without patches. Spiracles large and 
opened, spiracular plates brown. Marginal 
areas of body and some abdominal seg- 
ments behind siphunculi with distinct im- 
brications. Dorsal hairs of body thick and 
short, stout at apex. Head with 1 pair ce- 
phalic hairs, 2—3 pairs antennal tubercular 
hairs, 4 pairs dorsal hairs. Abdominal ter- 
gites I-VII each with 3 or 4 spinal, 1 pair 


895 


pleural and 4—8 pairs marginal hairs, tergite 
VIII with 2-3 pairs hairs. Length of ce- 
phalic hairs 0.04, length of marginal hairs 
on abdominal tergite I 0.03, and length of 
dorsal hairs on tergite VII 0.05, 0.61 times, 
0.49 times and 0.72 times as long as widest 
diameter of antennal segment III, respec- 
tively. Antenna 6-segmented, 1.20 times as 
long as body; length in proportion to seg- 
ments: Vi: 12.18; 100: 55.51 LO aa07, 
respectively. Antennal segment III with 29— 
36 hairs, length 0.03, 0.46 times as long as 
widest diameter of segment. Antennal seg- 
ment III with 46—73 large and small round 
secondary rhinaria, on entire segment. Ros- 
trum reaching midcoxae, ultimate rostral 
segment 1.40 times as long as second hind 
tarsal segment, with 8—9 pairs hairs, 5—6 
pairs accessory hairs among them. Hind fe- 
mur 1.21 times as long as antennal segment 
III. Hind tibia 0.79 times as long as body. 
Veins normal. Siphunculi swollen at distal 
half, constricted at apex, 0.23 times as long 
as body, 2.00 times as long as cauda. Cauda 
long, tapering, with 7 hairs. Anal plate with 
17-19 hairs. Other characters similar to ap- 
terous viviparous female. 

Alate male: Body long oval. Mounted 
specimens: Antenna, rostrum, distal half of 
femora, tibiae, tarsi, siphunculi, cauda, anal 
plate dark brown, other appendages slightly 
brown. Abdominal tergites I-V each with 1 
pair spino-pleural and 1 pair marginal 
patches. Dorsal hairs of body with sclerites 
at base. Dorsal hairs slightly short, ventral 
hairs longer than dorsal hairs. Antenna 6- 
segmented, 1.50 times as long as body. Sec- 
ondary rhinaria small round, on segments 
III-V; segment III with 53—63, segment IV 
with 8-11, segment V with 7-11. Fore- 
wing media two-branched, hindwing two 
obliques. Clasper shown in Fig. 13. Other 
characters similar to alate viviparous fe- 
male. 

Oviparous female: Body oval. Mounted 
specimens: Antennal segments I-III, distal 
part of segment IV, distal part of segment 
V, basal part of segment VI, distal part of 
rostrum, distal half of femora, tibiae, tarsi, 


896 


cauda, anal plate, distal / of siphunculi 
dark brown, other appendages slightly 
brown; basal Y; of siphunculi pale, rest of 
siphunculi slightly brown. Genital plate 
slightly brown. Antennal segment III with 
12-18 small round secondary rhinaria, on 
basal half of segment. Hind tibia swollen at 
basal half, with about 250 small round 
pseudo-sensoria. Other characters similar to 
apterous viviparous female. 

Diagnosis.—The new species differs 
from D. yezoense Miyazaki as follows: 
Clypeus with 4 hairs (yezoense, 2 hairs), 
basal and distal of antennal segment III and 
basal of segment IV pale (yezoense, black, 
except basal of segment III), and yellow or 
red in life (yezoense, green or bluish green). 
It differs from D. bogdouli Szelegiewicz as 
follows: Antennal segment V black (bog- 
douli, basal part of segment pale), yellow 
or red in life (bogdouli, shining brown in 
life), and length of dorsal hairs of body 0.33 
(bogdouli, 0.45). 

Etymology.—The species name is based 
on the host plant, Aconitum kusnezofii. 

Holotype.—Apterous viviparous female, 
No. 6804-1-2-3, 1978-VI-8, China, Beijing 
(Sanpu District, 116.4°E, 89.9°N), Col. G. 
X. Zhang and T. S. Zhong, on young twigs 
and upper of leaves of Aconitum kusnezofii 
Reichb. 

Paratypes.—Two apterous viviparous fe- 
males and 2 alate viviparous females, No. 
6804, other data same as holotype; 2 alate 
viviparous females, 5 apterous females and 
2 alate males, No. 6502, 1976-X-5, Col. B. 
L. Zhang, other data same as holotype; 2 
apterous viviparous females, | alate vivip- 
arous female, 3 alate males and 12 apterous 
females, No. 6628, 1977-X-14, other data 
same as holotype; 5 apterous viviparous fe- 
males and 3 alate viviparous females, No. 
6230, 1976-V-18, other data same as holo- 
type; 1 apterous viviparous female, 1 alate 
viviparous female and 2 apterous females, 
No. 11535, 1997-IX-11, China, Beijing 
(Xiaolongmen District, 116.0°E, 39.9°N), 
Col. J. G. Xiangyu, on Aconitum sp. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Delphiniobium gyamdaense Zhang 1981 
(Figs. 15-21) 


Delphiniobium gyamdaense Zhang 1981: 
264. 


Description.—Apterous viviparous fe- 
male: Body 3.04 in length, 1.50 in width. 
Body elliptical, yellow in life, siphunculi, 
antennae, legs and cauda black. Mounted 
specimens: Head and prothorax slightly 
brown, mesonotum, metanotum and abdo- 
men pale, without patches. Antennal seg- 
ments dark brown, except basal part of seg- 
ment III and distal part of segment VI; dis- 
tal part of rostrum, distal half of femora, 
distal part of tibiae and tarsi dark brown; 
distal ¥, of siphunculi, cauda and anal plate 
dark brown, others pale. Mesosternal furca 
with long stem. Median frontal tubercle 
slightly developed, antennal tubercles de- 
veloped, higher than median frontal tuber- 
cle. Antenna 6-segmented, about as long as 
body; length in proportion to segments I— 
VI: 13, 8, 100, 40, 40, 10+115. Antennal 
segment III with 44—46 round secondary 
rhinaria, over the entire segment. Length of 
hairs on segment III % as long as widest 
diameter of segment. Rostrum exceeding 
midcoxae, ultimate rostral segment 2.35 
times as long as its basal width, with 8 ac- 
cessory hairs. Siphunculi long barrel- 
shaped, swollen at middle, width at swollen 
part 1.90 times distal width, distal % with 
reticulations, 0.16 times as long as body, 
1.25 times as long as cauda. Cauda long 
tapering, indistinctly constricted, 2.56 times 
as long as its basal width, with 6 hairs. Anal 
plate with 16 hairs. 

Specimens examined.—Three apterous 
viviparous females, No. 11391, 1997-VI-8, 
China: Qinghai (Huzhu County, 101.9°E, 
36.8°N), Col. X. L. Chen, on Aconitum sp., 
Xizang (Gyamda County, 93.1°E, 29.9°N). 


Delphiniobium violisuctum Qiao and 
Zhang, new species 
(Figs. 22-31) 


Description.—Measurements: Apterous 
viviparous female: Body 3.13 in length, 


VOLUME 102, NUMBER 4 


897 


Figs. 15—21. 
segments I-IV. 17, Antennal segments V and VI. 18, Ultimate rostral segment. 19, Mesosternal furca. 20, 
Siphunculus. 21, Cauda. 


1.43 in width. Antenna 3.70, length of seg- 
ments I-VI 0.14, 0.10, 1.10, 0.56, 0.46, 
0.14+1.27, respectively. Ultimate rostral 
segment 0.12 in length. Hind femur 1.37, 
hind tibia 2.30, second hind tarsal segment 
0.16. Siphunculus 0.67 in length. Cauda 
0.45 in length. Alate viviparous female: 
Body 3.40 in length, 1.58 in width. 
Apterous viviparous female: Body large, 
dark green in life, on upper side of leaves. 
Mounted specimens: Head and prothorax 
brown, antennal segments I and IJ, distal 7% 
of segment III, distal % of segment IV, dis- 
tal half of segment V, basal part of segment 
VI and distal half of processus terminalis 
dark brown, others brown; apex of rostrum, 
distal ¥; of femora, distal ¥, of tibiae, tarsi, 


Delphiniobium gyamdaense, apterous viviparous female. 15, Dorsal view of head. 16, Antennal 


distal %4 of siphunculi, cauda and anal plate 
dark brown, coxae, trochanters, basal /; of 
femora, basal ”, of siphunculi and genital 
plate slightly brown, others brown. Spira- 
cles large, oval, opened; spiracular plate 
long oval, slightly pale. Mesosternal furca 
with long stem. Dorsal hairs of body thick, 
short, stout at apex. Head with 1 pair of 
cephalic hairs and 1| pair antennal tubercu- 
lar hairs, 4 pairs dorsal hairs; pronotum 
with 3 pairs spinal, 2 pairs pleural and 1 
pair marginal hairs; abdominal tergite I 
with 14 hairs, tergite VII with 4 hairs. 
Length of cephalic hairs 0.04, length of 
marginal hairs on tergite I 0.03, and length 
of dorsal hairs on tergite VII 0.05, 0.73 
times, 0.55 times and 0.91 times as long as 


898 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 22-31. 
Antennal segments I-IV. 24, Antennal segments V and VI. 25, Ultimate rostral segment. 26, Mesosternal furca. 
27, Dorsal view of abdominal tergites II—VIII. 28, Spiracle on abdominal segment II. 29, Siphunculus. 30, 
Cauda. 31, Alate viviparous female, antennal segment III. 


widest diameter of antennal segment III, re- 
spectively. Median frontal tubercle unde- 
veloped, antennal tubercles distinct, higher 
than median frontal tubercle, slightly di- 
verging at inner margins. Antenna 6-seg- 
mented, 1.18 times as long as body, length 
in proportion to segments I-VI: 14, 10, 100, 
54, 45, 14+123. Processus terminalis 9.11 
times as long as base of segment. Antennal 
hairs similar to dorsal hairs, antennal seg- 
ments I-VI each with 7, 4, 30, 11, 6, 3+3 
hairs, respectively, apex of preocessus ter- 
minalis with 3 hairs. Length of hairs on an- 
tennal segment III 0.04, 0.64 times as long 
as widest diameter of segment. Antennal 
segment III with 11 small round secondary 


Delphiniobium violisuctum. 22—30, Apterous viviparous female. 22, Dorsal view of head. 23, 


rhinaria, on basal half of segment. Rostrum 
reaching midcoxae, ultimate rostral seg- 
ment thick, 1.33 times as long as its basal 
width, 0.87 times as long as second hind 
tarsal segment; with 2 pairs accessory hairs. 
Legs normal. Hind femur 1.33 times as long 
as antennal segment III. Hind tibia 0.72 
times as long as body. Hairs on legs slightly 
longer than dorsal hairs of body. Length of 
hairs on hind tibia 0.07, 1.40 times as long 
as middle width of segment. First tarsal 
segment chaetotaxy: 3, 3, 3. Siphunculus 
long barrel-shaped, 3.42 times as long as 
basal width, 8.13 times as long as distal 
width, 0.21 times as long as body, 1.48 
times as long as cauda, swollen at middle, 


VOLUME 102, NUMBER 4 


distal ¥, of siphunculi with reticulations. 
Cauda long, tapering, slightly constricted at 
middle, 2.44 times as long as its basal 
width, with 7 hairs. Anal plate circular at 
apex, with 18 hairs. Genital plate with 15 
hairs, 2 long anterior hairs among them. 
Gonapophyses three, each with 5 or 6 hairs. 
Alate viviparous female: Body large. 
Mounted specimens: Basal of antennal seg- 
ment III pale, distal 4% of tibiae black, distal 
half of femora, and other antennal segments 
slightly brown, other appendages dark 
brown; dorsum of head brown; distal of 
rostrum, cauda, and anal plate greyish 
brown; basal ’/,; of siphunculi slightly 
brown, distal ¥; of siphunculi dark brown. 
Abdominal dorsum without patches. Anten- 
na 6-segmented, segment III with 42—47 
small round secondary rhinaria, on entire 
segment. Fore wing veins brown, media 
two-branched. Other characters similar to 
apterous viviparous females. 
Diagnosis.—The new species is similar 
to D. bogdouli Szelegiewicz from Mongolia 
by the median frontal tubercle and antennal 
tubercles, but differs as follows: antennal 
segment III with 42—47 secondary rhinaria 
in alatae (bogdouli, 58—62); ultimate rostral 
segment 0.87 times as long as second hind 
tarsal segment (bogdouli, longer than, 1.10 
times); siphunculi slightly swollen at mid- 
dle (bogdouli, indistinctly swollen); and 
dark green in life (bogdouli, shining 
brown). 
Etymology.—The new species is named 
based on the host plant, Viola verecunda. 
Holotype.—Apterous viviparous female, 
No. Y8070-1-1-1, China: Shanxi (Yangch- 
ene County; 112.4°E, 35.5°N), Col-.. Fei 
Zhao, 1996-VI-1, on Viola verecunda. 
Paratypes.—One alate viviparous female 
and 1 apterous larvae, No.Y8070, other 
data same as holotype. 


Delphiniobium yezoense Miyazaki 1971 


Delphiniobium yezoense Miyazaki 1971: 
34(1):40. 


Distribution.—No.Y4347, China: Hebei 
(Wulinshan Mountain, 117.4°E, 40.6°N), 


899 


1983-IX-12, Col. S.P. Tian; No. Y5718, Chi- 
na: Hebei (Kuancheng County, 118.4°E, 
40.6°N), 1983-IX-9, Col. S.P. Tian; 
No.Y5597, China: Hebei (Xiaowutai 
Mountain, 115.°E, 89.9°N), 1984-VI-22, 
Col. S.P. Tian; No.Y1598, China: Sichuan 
(An County, 104.4°E, 31.6°N), 1979-V-22, 
Col. H.Y. Li; No.Y1436, China: Sichuan 
(An County), 1978-IX, Col. H.Y. Li; Japan. 

Host-plants.—Aconitum kusnezoffi and 
A. comiichali (in China); A. chinense, A. ki- 
tadakense, A. yesoense and A. sachalinense 
(in Japan). 


ACKNOWLEDGMENTS 


We express our thanks to Mr. S. P. Tian, 
Mr. T. S. Zhong, Mr. H. Y. Li, Mrs. E Zhao, 
and Dr. X. L. Chen for collection of spec- 
imens and preparing some illustrations. The 
project was supported by the National Nat- 
ural Sciences Foundation of China (a grant: 
39700015) and a Biological and Technical 
Innovation Grant from the Chinese Acade- 
my of Sciences (A2999084). 


LITERATURE CITED 


Borner, C. 1950. Neue europdische Blattausarten. 
Selbsverlag. Naumbury, Private publication. 19 
PP- 

Hille Ris Lambers, D. 1947. Contributions to a Mono- 
graph of the Aphididae of Europe II. Temminckia 
VII: 179-319. 

Karsch, A. 1887. Myzus junackianus n. sp. Berliner 
Entomologica Zoologici 31: 21-23. 

Knechtel, W. K. and C. I. Manolache. 1944. Neue Blat- 
tlause fiir Rumanien. V. Academic Rumanine, 
Bulletin Section Science 25: 267-270. 

Mamontova-Solukha, V. O. 1966. Aphids of the So- 
vietic Karpats. Insects of Ukrainian Karpats and 
Transkarpattia, ser. Prolems of Zoological Akad 
Nauk Ukrainian RSR, Kiev, pp. 3—13. 

Miyazaki, M. 1971. A revision of the tribe Macrosi- 
phini of Japan (Homoptera: Aphididae, Aphidi- 
nae). Insecta Matsumurana 34(1): 1—9, 40—42. 

Mordvilko, A. K. 1914. [Insectes Hémiptéres (Insecta 
Hemiptera), Vol. I. Aphidoidea. Livraison I. (Fati- 
ne de la Russie et des pays limitrophes, fondée 
principalement sur les collections du Musée Zool- 
ogique de 1’Académie Impériale des Sciences de 
Petrograd.)] Petrograd. 236 pp. 

Paik, W. H. 1971. A new species of aphid from Mt. 
Hanla. Korean Society Plant Protection 10(2): 75— 
76. 


900 


. Insecta V. Illustrated Encyclopedia of Fauna 
& Flora of Korea 13: 245, 354—356. 

Remaudiére, G. and M. Remaudiére. 1997. Catalogue 
of the World’s Aphididae. Homoptera Aphidoidea. 
Institut National de la Recherche Agronomique. 
147, rue de I’ Universite, 75338 Paris Cedex 07: 
473 pp. 

Robinson, A. G. 1968. Two new species of aphids 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


(Homoptera: Aphididae) from Canada. The Ca- 
nadian Entomologist 100: 275—279. 

Szelegiewicz, H. 1969. Neue Blattlaus (Homoptera, 
Aphididae) aus der Mongolei. Annales Zoologici 
Warszawa 27: 169-194. 

Zhang, G. X. and T. S. Zhong. 1981. Homoptera: 
Aphidoidea, pp. 264—265. Jn Huang E S., ed., In- 
sects of Xizang, Beijing: Science Press, 600 pp. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 901-911 


NEW COSTA RICAN SPECIES OF PHYLLOPHAGA HARRIS 
(COLEOPTERA: MELOLONTHIDAE: MELOLONTHINAE) 


MIGUEL ANGEL MORON AND ANGEL SOLIS 


(MAM) Departamento de Entomologia, Instituto de Ecologia, A.C. (SEP-CONACYT). 
Apdo. Postal 63, Xalapa, Veracruz 91000, México (e-mail: moron.ma @ecologia.edu.mx); 
(AS) Departamento de Entomologia, Instituto Nacional de Biodiversidad (INBio), Santo 
Domingo de Heredia, Costa Rica 


Abstract.—Four new species of Phyllophaga are described from Costa Rica, Phyllo- 
phaga kohlmanniana and P. picadoana from the montane rain forest of the Monteverde 
Biosphere Reserve, Puntarenas, and P. janzeniana and P. guanacasteca from the tropical 
deciduous forests lowlands of Guanacaste. Drawings of male genital capsules, female 
genital plates, and tarsal claws are provided. 


Resuimen.—Se describen cuatro especies nuevas de Phyllophaga procedentes de tres 
localidades en Costa Rica: P. kohlmanniana y P. picadoana del bosque lluvioso de mon- 
tama de la Reserva de la Biosfera Monteverde, Puntarenas; P. janzeniana y P. guanacas- 
teca de los bosques tropicales caducifolios de las tierras bajas de Guanacaste. Se incluyen 
ilustraciones de las capsulas genitales masculinas, de las placas genitales femeninas y de 


las unas tarsales. 


Key Words: 


During curatorial work at the collection 
of INBio (1993-1999), we found specimens 
that represent four undescribed species of 
Phyllophaga from the provinces of Guan- 
acaste and Puntarenas, Costa Rica. Males of 
these species have deep cleft or bifid tarsal 
claws, while the females have dentate tarsal 
claws. Using the criteria of Saylor (1942), 
Sanderson (1958), or Mor6n (1986), the 
males of these species key to the subgenus 
Phytalus, whereas the females key to the 
subgenus Phyllophaga (s.str.). A similar 
situation is present with other species from 
Mexico, such as Phyllophaga ambigenus 
(Bates) and Phyllophaga mesophylla Mo- 
ron and Rivera (1992), both placed in the 
subgenus Phytalus based on the structure of 
the male tarsal claws and other characters. 
Awaiting for new advances in the supras- 
pecific classification of the genus, we place 


Phyllophaga, May beetles, new species, tropical forests, Costa Rica 


the presently described species as incerta 
sedis. In this paper, we describe males and 
females, address variation within the spe- 
cies, and present the precise distribution 
data for four new species of Phyllophaga 
from the montane rain forests and decidu- 
ous tropical forests in Costa Rica. 

Acronyms used in the text are as follows: 
CAS, California Academy of Sciences, San 
Francisco, USA; INBio, Instituto Nacional 
de Biodiversidad, Costa Rica; MXAL, pri- 
vate collection M.A. Moron, Xalapa, Méx- 
ico. 


Phyllophaga kohlmanniana Moron and 
Solis, new species 
(Figs. 1-5) 


Description.—Holotype male: Head and 
pronotum shiny dark brown, with dense 
vestiture of short whitish setae; elytra, ster- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


3 


Figs. 1—5. 


Phyllophaga kohlmanniana. 1, Male protarsal claw, lateral view. 2, Genital capsule, lateral view. 


3, Paramera, distal view. 4, Female protarsal claw, lateral view. 5, Female genital plates, ventral view. Scale 


lines = 1 mm, except figs. 1, 4 = 0.5 mm. 


nites and pygidium reddish dark brown, 
sligthly pruinose, completely covered with 
dense vestiture of uniform, whitish, short 
setae, that offer a macroscopic grayish, vel- 
vety appearance; mouth parts, tibiae and 
tarsi shiny reddish brown; coxae and fem- 
ora partially pruinose. Clypeus 3.7 * wider 
than long, anterior border curved, not sin- 
uate, with elevated margin, surface slightly 
concave, with many uniformly distributed, 
deep, round punctures each provided with 
short, erect seta. Frontoclypeal suture wide- 
ly sinuate and deeply impressed. Frons 1.6 
x wider than long, convex, finely punctate 
rugose, with short erect setae on entire sur- 
face. Antenna 10-segmented, with three 
segmented club, lamellae of 8th to 10th 
segments 1.6 X longer than length of pre- 
ceeding six segments combined; segments 
4 or 5 shorter than 3, with rounded promi- 
nences on anterior sides; segments 6 and 7 
wider than long, with semiconical promi- 
nences on anterior sides. Frons 3.7 X wider 


than dorsal diameter of eye. Eye canthi long 
and wide, with 13-14 setae. Labrum bi- 
lobed, widely sinuated, with scattered slen- 
der setae. Mentum slightly concave, im- 
punctate, with scarce slender setae, anterior 
border briefly sinuated, nearly straight. 
Pronotum 1.7 X wider than long and 2.9 X 
wider than frons. Pronotal disk shiny, but 
with basi-central area pruinose, total sur- 
face with deep, round punctures regularly 
separated by 1—2 diameters; lateral borders 
widely angulate, lateral marginal bead cren- 
ulate, with long, curved setae; basal bead 
complete, indicated by wide, deep sulcus; 
anterior angles straight, slightly prominent; 
posterior angles nearly straight, not promi- 
nent. Scutellum 1.5 X wider than long, with 
many small punctures. Elytron 2.3 X longer 
than wide, pruinose, densely and regularly 
punctate-setose; epipleural border narrow, 
extended along complete margin, slightly 
widened at % and ¥ of its length, provided 
with scarce slender setae at anterior third, 


VOLUME 102, NUMBER 4 


dull or pruinose at middle and posterior 
thirds; humeral calla rounded, prominent; 
apical calla rounded. Metathoracic wings 
completely developed. Propygidium prui- 
nose, with dense setiferous punctures. Py- 
gidium scarcely convex, pruinose, with 
round shaped, shallow rounded seitiferous 
punctures, regularly distributed; apical mar- 
gin with 16 long, slender setae; basal mar- 
gin effaced. Pterosterna with long, dense, 
yellowish setae. Visible abdominal sternites 
II to IV slightly depressed at midline, with 
a noticeable shallow sulcus; sternite V 
sligthly convex, pruinose, with dense seti- 
ferous punctures at middle; anal plate nar- 
rowed, transversely excavated, with elevat- 
ed anterior and posterior margins, and 10 
setae on apical border. Protibia nearly as 
long as protarsus (1:1.1), with external bor- 
der tridentate, preapical spur acute, straight, 
longer than 2nd protarsomerus (1.2:1). Me- 
sotibia with one oblique, well marked, se- 
tiferous carina on external side; upper api- 
cal spur straight, narrow, and 1.4 xX longer 
than lower spur. Metatibia slightly shorter 
than metatarsus (1:1.2), with one oblique 
setiferous carina on external side; upper 
apical spur articulated, straight, sharply 
pointed, as long as basal metatarsomere, 
and 1.5 X longer than lower spur; lower 
apical spur articulated, apex acute. Tarso- 
meres semicylindrical, elongate, with en- 
larged apex, some setae apically and two 
lines of setae along ventral side. Tarsal 
claws symmetrical, similar in all legs, deep- 
ly cleft, upper tooth shorter than lower 
tooth (Fig. 1). Genital capsule with short, 
narrowed parameres, dorsal and ventrally 
fused, ring shaped, apex with medium size, 
tooth-like projections (Fig. 3). Aedeagus 
very long, with preapical patches of spi- 
nules at sides and sclerotized dorsal support 
with sinuose flagellum (Fig. 2). Tectum (= 
phallobase) uniformly convex. Length of 
genital capsule from apex of parameres to 
border of basal piece: 6.8 mm. Total body 
length: 21.2 mm. Humeral width: 10.0 mm. 

Allotype female: Similar to the male ex- 
cept as follows: antenna with segments 5 


903 


and 6 fused, and appearing 9-segmented. 
Visible abdominal sternites II to VI convex, 
with dense setiferous punctures; anal plate 
1.8 X longer than male anal plate, very con- 
vex, punctate, with 12 slender setae at the 
apical border. Meso- and metatibiae each 
with one oblique, strong, setiferous carina 
near middle and other vague carina on basal 
third of external side. Both apical spurs of 
metatibia articulated, wide, lanceolate and 
curvated. Tarsal claws dentate, with long 
tooth near middle of ventral border (Fig. 4). 
Ventral genital plates well sclerotized, near- 
ly symmetrical, elongated, with short setae 
at distal process; dorsal genital plates fused 
with ventral ones toward lateral sides, with 
acure apices directed mesad and setae on 
distal borders (Fig. 5). Total body length: 
20.6 mm. Humeral width: 9.8 mm. 

Variation.—Male: Similar to holotype 
except as follows: total body length: 19.8— 
21.5 mm, humeral width: 9.6—10.3 mm, 
pronotum of some specimens more reddish 
than holotype, other specimens with anten- 
nal club as long as preceeding six segments 
combined. Female similar to allotype ex- 
cept as follows: pygidium with more seti- 
ferous punctures; total body length: 19.5— 
20.4 mm; humeral width: 9.6—9.8 mm. 

Type material.—Described from 5 6, 4 
?. Holotype 6 INBio: Costa Rica: Puntar- 
enas, Monteverde, La Casona, IV-92, 1,520 
m, N. Obando. Allotype 2 INBio: Costa 
Rica: Guanacaste, SW Volcan Cacao, Est. 
Mengo, II-89, 1,100 m, GNP Biodiversity 
Survey. Paratypes INBio, MXAL: same 
data as holotype (2 ¢ and 2 ¢); same data 
as allotype (2 6, 1 @). 

Type locality.—La Casona, Estaci6n 
Monteverde, province of Puntarenas, Costa 
Rica (aprox. 10°30’N; 85°10’ W). 

Biological data.—Specimens of P. kohl- 
manniana were collected at lights near 
cloud forests and tropical premontane for- 
ests located between 1,100—1,520 m. Phe- 
nology: February (4), April (5). Other spe- 
cies of Phyllophaga flying at the same time 
were P. (Phyllophaga) tapantina Mor6n 


904 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


8 


Figs. 6-11. 


Phyllophaga janzeniana. 6, Male protarsal claw, lateral view. 7, Genital capsule, lateral view. 


8, Paramera, distal view. 9, Paramera, ventro-apical view. 10, Female tarsal claw, lateral view. 11, Female genital 
plates, ventral view. Scale lines = 1 mm, except figs. 6, 10, 11 = 0.5 mm. 


and Solis, P. (Phyllophaga) tilarana Mor6n 
and Solis and P. picadoana, n.sp. 
Remarks.—Phyllophaga kohlmanniana 
does not belong to any species group de- 
scribed by Moron (1986). The structure of 
the male tarsal claws suggests placement in 
the subgenus Phytalus, but the form of the 
female tarsal claws indicates placement in 
the subgenus Phyllophaga (s.str.), as is 
known for P. ambygenus (Bates) and some 
other Mexican and Central American spe- 
cies. By the color, vestiture, body size, 
length of tarsi, and form of clypeus is sim- 
ilar to P. (Phytalus) zeteki Saylor, 1942, de- 
scribed from Barro Colorado Island, Pana- 
ma, but the shape and details of the male 
genital capsule (Fig. 2), sculpture and ves- 


titure of the sternites, shape of tarsal claws 
(Fig. 1) and proportions of antennal seg- 
ments will aid in the recognition of this new 
species. 

Etymology.—This new species is dedi- 
cated to our friend, Dr. Bert Kohlmann, 
whose enthusiastic devotion to the study of 
the scarab beetles of Mexico and Central 
America is well known to coleopterists and 
also to people interested in insect ecology 
and biogeography. 


Phyllophaga janzeniana Moron and 
Solis, new species 
(Figs. 6-11) 


Description.—Holotype male: Clypeus, 
frons and pronotum shiny dark reddish 


VOLUME 102, NUMBER 4 


brown; elytra and pygidium shiny reddish 
brown, without macroscopic vestiture; 
mouth parts, sterna and legs shiny testa- 
ceous reddish brown. Clypeus 3.6 X wider 
than long, anterior border rounded, not sin- 
uate, with elevated margin, surface slightly 
concave, with some wide, deep, coarse, 
round punctures without microscopic setae. 
Frontoclypeal suture sinuate and vaguely 
impressed. Frons 2.0 X wider than long, 
convex, coarsely punctate rugose, with 
transverse keel on anterior half and some 
medium size setae at sides and minute setae 
scattered on the posterior half of disk. An- 
tenna 9-segmented, with 3 segmented club, 
lamellae as long as the length of preceeding 
5 segments combined; 5th segment longer 
than 4 or 3; segment 6 compresed with 
semiconical anterior process. Frons 3.3 X 
wider than dorsal diameter of eye. Eye can- 
thi long and wide, with 9—10 setae. Labrum 
widely concave, sligthly bilobed, with scat- 
tered slender setae. Mentum slightly con- 
cave, impunctate, with scarce, slender setae, 
anterior border briefly notched. Pronotum 
1.6 X wider than long and 2.9 X wider than 
frons. Pronotal disk shiny, with deep, 
coarse umbiliform punctures irregularly 
separated by 1-3 diameters; lateral borders 
widely angulate, lateral marginal bead 
widely crenulated, with scattered, long, 
slender setae; basal bead vaguely indicated 
by punctures, mainly toward the sides; an- 
terior angles straight, prominent; posterior 
angles widely obtuse, rounded. Scutellum 
1.3 X wider than long, with some irregular 
punctures. Elytron 2.8 < longer than wider, 
shiny, densely rugo-punctate; epipleural 
border very narrow, extended along com- 
plete margin, provided with some scattered 
short setae; humeral calla rounded, promi- 
nent; apical calla rounded. Metathoracic 
wings completely developed. Propygidium 
glabrous, with distal half nearly shiny and 
basal half slightly pruinose, with sparse um- 
biliform punctures. Pygidium moderately 
convex, shiny, glabrous, coarsely rugo- 
punctate; apical margin with 14 long, slen- 
der setae; basal margin effaced medially. 


905 


Pterosterna with medium size, moderately 
dense, yellowish setae. Visible abdominal 
sternites II and IV slightly convex, nearly 
polished and glabrous toward the middline; 
sternite V convex, shiny, irregularly rugo- 
punctate with scarce, scattered setae at mid- 
dle; anal plate narrowed, transversely con- 
cave, irregularly punctate, anterior and pos- 
terior borders elevated, with some slender 
setae. Protibia slightly shorter than protar- 
sus (1:1.2), with external border tridentate, 
preapical spur acute, nearly straight, as long 
as 2nd protarsomerus. Mesotibia with one 
oblique, sharp setiferous carina on external 
side; upper apical spur straight, narrow, and 
1.1 X longer than lower spur. Metatibia 
nearly as long as metatarsus, with one 
oblique, sharp setiferous carina on external 
side; upper apical spur articulated, angulat- 
ed, sharply pointed, 1.2 < longer than basal 
metatarsomere, and 1.1 X longer than lower 
spur; lower apical spur articulated, with 
acute apex. Tarsomeres semicylindrical, 
elongate, with enlarged apex, some setae 
apically and two lines of setae on ventral 
side. Tarsal claws symmetrical, similar in 
all legs, widely and deeply cleft, upper 
tooth nearly as long as lower tooth, poste- 
rior ventral border slightly serrated (Fig. 6). 
Genital capsule with medium size paramer- 
es, dorsally fused, elongated with apex ex- 
panded and curvated toward the middle line 
(Figs. 7-8); ventrally the paramera are not 
fused (Fig. 9). Aedeagus long, with a preap- 
ical dorsal sclerotized plate that support a 
pair of long curved spine-like sclerotized 
structures (Fig. 7). Tectum uniformly con- 
vex. Length of genital capsule from apex of 
parameres to border of basal piece: 3.7 mm. 
Total body length: 11.9 mm. Humeral 
width: 5.0 mm. 

Allotype female: Similar to the male ex- 
cept as follows: anterior angles of clypeus 
more wide and rounded, expanded; anten- 
nal club as long as the length of four pre- 
ceeding segments; pronotum with anterior 
angles more prominent and lateral bead no- 
ticeably crenulate. Visible abdominal ster- 
nites II to IV convex, with scattered seti- 


906 


ferous punctures; sternite V more long and 
convex; anal plate 2 X< longer than male 
anal plate, convex, punctate, with 10 slen- 
der setae near the posterior border. Pygidi- 
um with central part of disk slightly flat- 
tened. Both apical spurs of metatibia slight- 
ly curvated with the apex rounded. Tarsal 
claws with upper tooth slightly shorter than 
ventral one (Fig. 10). Ventral genital plates 
with basal half well sclerotized and distal 
part membranous, sclerotized only at bor- 
ders, sligthly asymmetrical, basally round- 
ed, elongated to apex, with some setae on 
border; dorsal genital plates poorly sclero- 
tized, fused medially, with some setae at 
each side (Fig. 11). Total body length: 11.2 
mm. Humeral width: 5.2 mm. 

Paratype variation.—Male similar to ho- 
lotype except in total body length: 10.8— 
11.8 mm, humeral width: 4.8—5.2 mm, 
pronotum and elytra of some specimens 
darker than holotype, other specimens with 
frontal transverse carina more or less ac- 
centuated. Female similar to allotype except 
as follows: pronotum, elytra and pygidium 
with more or less punctures; total body 
length: 10.8—11.5 mm; humeral width: 5.0— 
5.4 mm. 

Type material.—Described from 8 6, 10 
?. Holotype 6 INBio: Costa Rica: Guan- 
acaste, 12 km SE La Cruz, Cerro El Hacha, 
Casa Oeste, VI-87, 300 m, GNP Biodiver- 
sity Survey. Allotype 2°: same data as ho- 
lotype (INBio). Paratypes CAS, INBio, 
MXAL: same data as holotype (4 6, 1 2); 
Guanacaste, PN. Rincon de La Vieja, Est. 
Las Pailas, 1/22-VII-92, 800 m, D. Garcia 
GB. #8-2)); 

Type locality—Cerro El Hacha, 12 km 
SE La Cruz, province of Guanacaste, Costa 
Rica (approx. 10°58'N; 85°32'W). 

Biological data.x—Specimens of P. jan- 
zeniana were collected at lights in tropical 
deciduous forests located between 300—800 
m. Phenology: June (7), July (11). Other 
species of Phyllophaga flying at the same 
time were P. (Chlaenobia) scabripyga 
(Bates), P. (Phytalus) obsoleta (Blanchard), 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


P. (Phyllophaga) elenans Saylor and P. 
(s.str.) hondurasana Moser. 

Remarks.—Phyllophaga janzeniana does 
not belong to any species group proposed 
by Mor6én (1986). It may be related to P. 
soctona Mor6n from Chiapas, México, 
which is also a small species with frontal 
transverse carina, 9-segmented antenna and 
similar tarsal claws, but the shape and de- 
tails of the male genital capsule (Figs. 7—8) 
and female genital plates (Fig. 11) are very 
different. Each paramere of P. soctona have 
two large, apical teeth, and dorsal genital 
plates of female are not fused medially. 

Etymology.—This new species is dedi- 
cated to Dr. Daniel H. Janzen, whose inter- 
est in the study and conservation of tropical 
forests, mainly in the Guanacaste area, is 
well known. 


Phyllophaga picadoana Moré6n and 
Solis, new species 
(Figs. 12-16) 


Description.—Holotype male: Head, 
pronotum and pygidium shiny dark reddish 
brown; with sparce vestiture of short to 
long setae; mouth parts, sterna and legs 
shiny reddish brown. Clypeus 3.2 * wider 
than long, anterior border shallowly and 
widely sinuated, with margin elevated, sur- 
face nearly flat, with large, shallow, round 
punctures and some scattered short setae. 
Frontoclypeal suture fine, vaguely indicated 
by a wide, shallow sinuate sulcus. Frons 1.7 
x wider than long, convex, coarsely punc- 
tate rugose, covered with short to medium 
size slender setae. Antenna 10-segmented, 
with 3-segmented club, lamellae 2 < longer 
than length of preceeding 5 segments com- 
bined. Frons 4.1 X wider than dorsal di- 
ameter of eye. Eye canthi long and wide, 
with 7-8 setae. Labrum widely sinuate, 
slightly bilobed, with scattered slender se- 
tae. Mentum slightly convex, impunctate, 
with few slender setae, anterior border shal- 
lowly and widely sinuated. Pronotum 1.7 X 
wider than long and 3 X wider than frons. 
Pronotal disk shiny, with shallow, round 
shaped, large setiferous punctures irregular- 


VOLUME 102, NUMBER 4 


907 


16 


Figs. 12-16. Phyllophaga picadoana. 12, Male protarsal claw, lateral view. 13, Genital capsule, lateral view. 
14, Paramera, distal view. 15, Female protarsal claw, lateral view. 16, Female genital plates, ventral view. Scale 


lines = | mm, except figs. 12, 15 = 0.5 mm. 


ly separated from one another by 1-3 di- 
ameters; lateral borders widely angulate, 
lateral marginal bead slightly crenulated, 
mainly toward basal half, with scattered, 
long, slender setae; basal bead erased, only 
with shallow sulcus, mainly toward sides; 
anterior angles acute, prominent; posterior 
angles slightly obtuse, not prominent. Scu- 
tellum 1.6 X wider than long, with 10 
rounded punctures. Elytron 2.4 X longer 
than wide, densely punctate-rugose, mostly 
shiny, except the preapical area and distal 
half of the lateral borders that have vague 
pruinose vestiture; epipleural border mod- 
erately narrowed, extended along complete 
margin, provided with scattered short, slen- 
der setae; humeral calla rounded, promi- 
nent; apical calla rounded. Metathoracic 
wings completely developed. Propygidium 
shiny, with dense setiferous punctures. Py- 
gidium moderately convex, shiny, rugose, 
with round, shallow setiferous punctures ir- 
regularly distributed; apical margin with 18 


long, slender setae; basal margin effaced 
medially. Pterosterna with long, dense, yel- 
lowish setae. Visible abdominal sternites II 
and IV slightly depressed, polished and 
nearly glabrous at the middline; sternite V 
convex, with a patch of granules and erect 
short setae on middle and with a wide, shal- 
low transverse sulcus before posterior bor- 
der; anal plate sligthly concave, vaguely 
furrowed at middle with some erect, medi- 
um size setae and anterior and posterior 
borders slightly elevated. Protibia slightly 
shorter than protarsus (1:1.2), with external 
border tridentate, preapical spur acute, near- 
ly straight, slightly longer than 2nd protar- 
somerus (1:1.2). Mesotibia with one 
oblique, sharp, setiferous carina on external 
side; upper apical spur straight, narrowed, 
as long as the lower spur. Metatibia slightly 
shorter than metatarsus (1: 1.2), with one 
oblique, sharp, setiferous carina on external 
side; upper apical spur nearly lanceolate, 
curved, with rounded apex, as long as basal 


908 


metatarsomere, and 1.4 X longer than lower 
spur; lower apical spur articulated, with 
rounded apex. Tarsomeres semicylindrical, 
elongate, with enlarged apex, some setae 
apically and two rows of setae with a fine 
longitudinal keel in middle of ventral side. 
Protarsomeres 1—3 with subapical short 
spines. Tarsal claws symmetrical, similar in 
all legs, deep cleft, upper tooth as long as 
lower tooth, with middle of ventral border 
finely serrate (Fig. 12). Genital capsule with 
proportionately short parameres, dorsally 
fused, apical thirth canaliculated by inner 
side (Fig. 14). Aedeagus large, with wide 
apical patches of spinules and dorsal preap- 
ical tuft of strong setae on a wide sclero- 
tized support (Fig. 13). Tectum uniformly 
convex. Length of genital capsule from 
apex of parameres to border of basal piece: 
4.1 mm. Total body length: 17.1 mm. Hu- 
meral width: 7.8 mm. 

Allotype female: Similar to male except 
as follows: elytra with long erect setae near 
scutellum and toward apex. Pygidium less 
convex and rugose. Visible abdominal ster- 
nites II to V convex, with scattered setifer- 
ous punctures; anal plate convex, punctate, 
with many slender setae. Apical spurs of 
metatibia wide, lanceolate and curved. Tar- 
sal claws with preapical tooth on ventral 
border, nearly as long as apical tooth, with 
part of ventral border serrate (Fig. 15). Ven- 
tral genital plates slightly sclerotized, nearly 
symmetrical, with apical border rounded 
(Fig. 16). Total body length: 17.3 mm. Hu- 
meral width: 8.1 mm. 

Paratype variation.—Male similar to ho- 
lotype except in total body length: 17.0— 
17.4 mm, humeral width: 7.6—7.9 mm, dor- 
sal vestiture of some specimens less dense 
than holotype, other specimens with anten- 
nal club 1.8—2.1 X longer than length of 
preceeding five segments combined. Spec- 
imens from Chirripo have a darker color, 
distal half of paramera less canaliculated 
and have a wide shallow concavity on the 
dorsal part of parameral base. Female sim- 
ilar to allotype except as follows: elytra and 
pygidium with more or less setiferous punc- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


tures; total body length: 16.8—17.3 mm; hu- 
meral width: 7.9—8.1 mm. 

Type material—Described from 10 6 
and 8 2. Holotype ¢ INBio: Costa Rica: 
Puntarenas, R.B. Monteverde, La Casona, 
I-92, 1,520 m, N. Obando. Allotype 2 IN- 
Bio: same data as holotype except III-1994. 
Paratypes CAS, INBio, MXAL: same data 
as holotype (5 ¢); same data as allotype (2 
3,3 2); Cartago, Turrialba, Chirripo, Gra- 
no de Oro, I[X-92, 1,120 m, P. Campos (2 
6a4Ae®): 

Type locality—La Casona, Reserva Biol- 
6gica Monteverde, province of Puntarenas, 
Costa Rica (approx. 10°30'N; 85°10'W). 

Biological data.—Specimens of P. pica- 
doana were collected at lights near montane 
tropical forests and coffee plantations lo- 
cated at 1,120—1,520 m. Phenology: March 
(12), September (6). Other species of Phyl- 
lophaga flying at the same time were P. 
(Phyllophaga) tilarana Moron and Solis, P. 
(s.str.) tapantina Moroén and Solis and P. 
kohlmanniana Moron and Solis. 

Remarks.—Phyllophaga picadoana does 
not belong to any species group proposed 
by Moron (1986). Externally this species 
resembles P. (Phyllophaga) brevidens 
Bates and allies, but the male and female 
tarsal claws of P. brevidens have a small 
tooth at the middle of ventral border, each 
paramere is deeply bifurcate and the aedea- 
gus do not present wide apical patches of 
spinules. 

Etymology.—The name of this new spe- 
cies is dedicated to the memory of the for- 
mer Costa Rican biologist, Dr. Clodomiro 
Picado Twight, who devoted much of his 
life to the study of animals, tropical ecology 
and natural richess of that country. 


Phyllophaga guanacasteca Moron and 
Solis, new species 
(Figs. 17-22) 


Description.—Holotype male: Head, 
pronotum and elytra dark reddish brown 
with dense pruinose greyish vestiture; 
mouth parts, pygidium and legs shiny red- 
dish brown; sternites light reddish brown 


VOLUME 102, NUMBER 4 


909 


Figs. 17-22. 


Phyllophaga guanacasteca. 17, Male protarsal claw, lateral view. 18, Male hind tarsal claw, 


lateral view. 19, Genital capsule, lateral view. 20, Paramera, distal view. 21, Female protarsal claw, lateral view. 


22. Female genital plates, ventral view. Scale lines = 


with fine pruinose vestiture. Clypeus 3.4 X 
wider than long, anterior border wide and 
deeply sinuate, with elevated margin, sur- 
face slightly convex, with many uniformly 
distributed, shallow large punctures, with- 
out setae. Frontoclypeal suture sinuate and 
deeply impressed. Frons 1.4 < wider than 
long, convex, densely punctate, without se- 
tae. Antenna 10-segmented, with 3-seg- 
mented club, lamellae 1.3 xX longer than 
length of preceeding 6 segments combined. 


1 mm, except figs. 17, 18, 21 = 0.5 mm. 


Frons 2.5 X wider than dorsal diameter of 
eye. Eye canthi long and wide, with 9-11 
setae. Labrum bilobed, widely sinuated, 
with scattered slender setae. Mentum slight- 
ly convex, impunctate, with few slender se- 
tae, anterior border nearly straight. Prono- 
tum 1.6 X wider than long and 2.9 X wider 
than frons. Pronotal disk pruinose, with 
shallow, round small punctures regularly 
separated from one another by 2—4 diame- 
ters; lateral borders widely rounded, lateral 


910 


marginal bead entire, with 2-3 scattered 
slender setae; basal bead indicated by punc- 
tures, mainly toward sides, where shallow 
sulcus are also indicated; anterior angles 
straight, slightly prominent; posterior an- 
gles obtuse, slightly prominent. Scutellum 
1.5 X wider than long, scattered small 
punctures. Elytron 2.6 X longer than wide, 
pruinose, uniformly punctate with some in- 
terstriae slightly elevated; epipleural border 
narrow, extended along complete margin, 
provided with some scattered short setae; 
humeral calla rounded, prominent; apical 
calla rounded. Metathoracic wings com- 
pletely developed. Propygidium pruinose, 
with uniform round, small punctures. Py- 
gidium convex, shiny, with round, shallow 
punctures irregularly distributed; apical 
margin with 14 long, slender setae; basal 
margin effaced medially. Pterosterna with 
medium size, moderately dense, yellowish 
setae. Visible abdominal sternites II and III 
slightly depressed, pruinose, with small, 
black, granules at midline; sternite [V with 
dentiform irregular black granules near pos- 
terior border; sternite V with rounded area 
scarcely prominent at middle, covered with 
fine rugosities, granules and short setae; 
anal plate narrowed, sligthly convex, with 
some short setae near midline and 6 slender 
setae on apical border. Protibia shorter than 
protarsus (1:1.8), with external border tri- 
dentate, but basal tooth very reduced; 
preapical spur acute, nearly straight, shorter 
than 2nd protarsomerus (1:2.2). Mesotibia 
with one oblique, setiferous carina on ex- 
ternal side; upper apical spur straight, nar- 
rowed to the apex, and 1.8 X longer than 
lower spur; lower spur with truncated apex. 
Metatibia shorter than metatarsus (1:1:3), 
with one oblique, setiferous carina on ex- 
ternal side; upper apical spur straight, 
sharply pointed, slightly longer than basal 
metatarsomere (1.2:1), and 1.9 X longer 
than lower spur; lower apical spur articu- 
lated, curved, with rounded apex. Protar- 
someres 1—4 slightly concave by ventral 
sides, with lateral rows of setae. Meso- and 
metatarsomeres semicylindrical, elongate, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


with enlarged apex, some setae apically and 
two lines of setae on ventral side. Protarsal 
claws slightly asymmetrical, deeply cleft, 
upper tooth longer and narrower than lower 
tooth (Fig. 17). Meso- and metatarsal claws 
symmetrical, deeply cleft, with upper tooth 
slightly longer than lower tooth (Fig. 18). 
Genital capsule with long parameres, dor- 
sally fused, with preapical tooth-like pro- 
jections on the external and internal borders 
(Figs. 19-20). Aedeagus short, with preap- 
ical, large, sclerotized asymmetrical plate 
(Fig. 19). Tectum with two large latero-dis- 
tal rounded prominences (Fig. 19). Length 
of genital capsule from apex of parameres 
to border of basal piece: 5.9 mm. Total 
body length: 19.1 mm. Humeral width: 7.9 
mm. 

Allotype female: Similar to male except 
as follows: clypeal disk with more deep 
punctures; antennal club shorter than length 
of five preceeding segments (1:1.3). Pygid- 
ium smaller, slightly convex, with more 
punctures, mainly toward apex. Visible ab- 
dominal sternites II to TV convex, with scat- 
tered setae; sternite V slightly longer than 
the IV segment, with more setae; anal plate 
convex, punctate, with 30 slender setae. 
Meso- and metatibiae each with one 
oblique, strong, setiferous carina on exter- 
nal side. Both apical spurs of metatibia ar- 
ticulated, wide, lanceolate and curved. Pro- 
tarsal segments 1—4 semicylindrical, elon- 
gated with enlarged apex, but not concave 
on ventral surface. Tarsal claws clearly den- 
tate, with apical tooth nearly as long as pos- 
terior one (Fig. 21). Ventral genital plates 
well sclerotized, symmetrical, rounded with 
wide apical borders; dorsal genital plates 
ovate, with slightly projected rounded apex 
(Fig. 22). Total body length: 20.0 mm. Hu- 
meral width: 8.4 mm. 

Paratype variation.—Male similar to ho- 
lotype except in total body length: 18.6— 
19.3 mm, humeral width: 7.6—8.0 mm, dor- 
sal color of some specimens darker than ho- 
lotype or with the pruinose vestiture more 
or less dense. Female similar to allotype ex- 
cept as follows: frons and pygidium with 


VOLUME 102, NUMBER 4 


more punctures; total body length: 19.5— 
20.3 mm; humeral width: 8.3—8.4 mm. 

Type series.—Described from 4 d, 3 9. 
Holotype ¢ INBio: Costa Rica: Estacion 
Pitilla, 9 km S Sta. Cecilia, PN. Guana- 
caste, Guanacaste, Costa Rica, V-94, 700 
m, C. Moraga. Allotype 2 INBio: same 
data except III/IV-93, Malaise. Paratypes 
INBio, MXAL: same data as holotype (3 6, 
ey): 

Type locality—Estacion Pitilla, 9 km S 
Santa Cecilia, Guanacaste National Park, 
province of Guanacaste, Costa Rica 
(approx. 10°59'26"N; 85°25'40"W). 

Biological data——Males and females of 
P. guanacasteca were collected at lights 
near deciduous tropical forest located at 
700 m. Phenology: March—April (1), May 
(6). Other species of Phyllophaga flying at 
the same time were P. hondurasana Moser 
and P. guapiles Saylot. 

Remarks.—Phyllophaga guanacasteca 
does not belong to any species group de- 
fined by Moron (1986). It is general ap- 
pearance suggest a relationship with P. 
(Phytalus) pruinosa (Blanchard), but the ar- 
ticulated lower metatibial spur in the male, 
and the dentate tarsal claws of the female 
do not match with the diagnostic characters 
of the group pruinosa. The shape and de- 
tails of the male genital capsule (Figs. 19— 
20) and female genital plates (Fig. 22), 
length of lower metatibial spur of male, 
sculpture of the pygidium and sternites and 
shape of protarsal claws (Fig. 17) will aid 
in the recognition of this new species. 

Etymology.—Specific epithet derived 


OV 


from the name of the province of Guana- 
caste, to which this species appears to be 
restricted. 


ACKNOWLEDGMENTS 


Study of the collections of INBio made 
by Miguel A. Moron was made possible by 
the support of Instituto de Ecologia, Xalapa 
and Programa Diversidad Biolégica, CYT- 
ED-México (1993), and by the support of 
INBio (1998). Research done at Instituto de 
Ecologia, Xalapa by Angel Solis was made 
possible with the financial aid of INBio 
(1997) and CONACYT (1999). We appre- 
ciate corrections and suggestions of two 
anonymous reviewers for better English ex- 
pression. This paper is a contribution to the 
project ‘““Sistematica y Biologia del género 
Phyllophaga en México y América Cen- 
tral’? (225260-25723-N CONACYT Méxi- 
CO). 


LITERATURE CITED 


Moron, M. A. 1986. El género Phyllophaga en Méx- 
ico. Morfologia, Distribuci6n y Sistemdatica Su- 
praespecifica (Insecta: Coleoptera). Publ. 20, In- 
stituto de Ecologia, México, 341 pp. 

Moron M. A. and L. E. Rivera-Cervantes. 1992. Dos 
especies nuevas de Phyllophaga (Phytalus) (Co- 
leoptera: Melolonthidae) de la Sierra de Manan- 
tlan, Jalisco, México. Anales Instituto de Biologia 
Universidad Nacional Autonoma de México, serie 
Zoologia 63(1): 79-87. 

Sanderson, M. W. 1958. Faunal affinities of Arizona 
Phyllophaga, with notes and descriptions of new 
species. Journal of the Kansas Entomological So- 
ciety 31: 158-173. 

Saylor, L. W. 1942. Notes on beetles related to Phyl- 
lophaga Harris with descriptions on new genera 
and subgenera. Proceedings of the United States 
National Museum 92(3145): 157-165. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 912-918 


TWO ADDITIONAL SPECIES OF ROBBER FLIES OF THE GENUS 
OMMATIUS WIEDEMANN (DIPTERA: ASILIDAE) FROM THE BAHAMAS 
AND WITH REPLACEMENT NAMES FOR TWO OTHER SPECIES 


A. G. SCARBROUGH 


Department of Biology, Towson University, Baltimore, MD 21252, U.S.A. (e-mail: 


ascarbrough @ towson.edu) 


Abstract.—Two species of robber flies of the genus Ommatius Wiedemann are reported 
from the Bahamas, increasing the number of species to six. Ommatius mariae, Nn. sp., 
from Eleuthera and Nassau, is described, illustrated, and contrasted with O. hanebrinki 
Scarbrough and Rutkauskas. Ommatius mariae, O. hanebrinki, and O. hispidus Scar- 
brough from the hanebrinki species group. A Cuban species, O. lineolatus Scarbrough is 
reported for the first time from Long Island in the Bahamas. Two new replacement names 
are proposed: Ommatius dignus new name for Ommatius dimidiatus Scarbrough and 
Ommatius fimbrillus new name for Ommatius fimbriatus Scarbrough and Poinar. 


Key Words: 


The robber flies of the genus Ommatius 
Wiedemann from the Bahama Islands are 
poorly known. To date four species are re- 
ported from single localities, 1.e., O. abana 
Curran (1953) from Bimini, O. hanebrinki 
Scarbrough and Rutkauskas (1983) from 
San Salvador, O. membranous Scarbrough 
(1985a) from Rum Cay, and _ setiferous 
Scarbrough (1988) from Mayaguana. Fur- 
thermore, only one sex is known for all spe- 
cies except for O. hanebrinki. This paper 
reports a new species of Ommatius Wie- 
demann from Eleuthera and Nassau, and the 
discovery of a Cuban species, O. lineolatus 
Scarbrough (1988), from Long Island. Two 
replacement names are proposed to correct 
nomenclatural errors. 


METHODS 


General methods and terminology follow 
that described by Scarbrough (1997) and 
McAlpine (1981), respectively. Ratios used 
in the text are as follows: face:head width 


nomenclature, Asilidae, Ommatius, new species, new records, Bahamas 


ratio (FHWR) is the greatest width of the 
head in front profile divided by the width 
of the face at the base of the antennae; cell 
m1 width ratio (m1WR) is the ratio of the 
width of the cell at the basal third and api- 
cal two-thirds divided by the width of the 
base; hind femoral width:length ratio 
(HFWLR) is the greatest dorsal length di- 
vided by the greatest width. 

Acronyms of museums used in the text 
are as follows: TUMZ, Museum of Zoolo- 
gy, Towson University, Baltimore, MD; 
AMNH, American Museum of Natural His- 
tory, NY; CMNH, Carnegie Museum of 
Natural History, Pittsburgh, PA; USNM, 
National Museum of Natural History, 
Smithsonian Institution, Washington, DC; 
FSCA, Florida State Collection of Arthro- 
pods, Gainesville, FL; GPAC, George Poin- 
ar Collection of Amber, Department of En- 
tomology, Oregon State University, Cor- 
vallis, OR. 


VOLUME 102, NUMBER 4 


Ommatius mariae Scarbrough, 
new species 
(Figs. 1-9) 


Male.—Brown. Body 11.1—12.3 mm; 
wing 8.3-8.6 mm. Head: Dull yellowish 
gray tomentose; frons, vertex, and narrow 
margin of occiput dorsally more yellow; 
vestiture largely whitish to pale yellow; 
FHWR 1.0:6.3—1.0:6.9. Antenna mostly 
brown setose, whitish setae present below 
scape; flagellum broadly oval, slightly lon- 
ger than wide, and slightly longer than 
scape. Frons with margins parallel. Occiput 
with 3-5 short, thick, brown postocular 
bristles; bristles slightly curved forward, 
apices of bristles above eye. 

Thorax: Pronotum yellowish gray tomen- 
tose with 2 brown and 2 whitish bristles. 
Mesonotum with wide median stripe divid- 
ed medially by a thin, light yellowish to- 
mentose line; 2 lateral spots brown tomen- 
tose spots present; tomentum otherwise yel- 
low to yellowish gray; setae sparse, limited 
to lateral margins; 3 posterior dorsocentral 
and 4 lateral stout, dark brown bristles. Scu- 
tellum yellowish tomentose with scattered 
whitish setae; 2 short, brown, marginal se- 
tae sometimes present, each about half as 
long as dorsal setae; preapical groove ab- 
sent. Pleura dull yellowish gray tomentose 
anteriorly, more grayish posteriorly; vesti- 
ture whitish; halter yellow. 

Wing: Margin anteriorly straight, without 
a costal bulge. Cell r4 narrow and long, 
sides only slightly divergent apically; base 
before apex of cell d. Crossvein r-m just 
beyond middle of cell d, slightly shorter 
than or as long as vein CuA1+M3. Cell m1 
Vass 120:1.9:1-4-1.0:2.7:2.1. 

Legs: Coxae brown with grayish tomen- 
tum and pale yellow to white vestiture; fore 
coxa with numerous stout bristles anterior- 
ly. Trochanters brown. Femora mostly yel- 
low with yellowish bristles; fore femur 
mostly brown anteriorly with a narrow yel- 
low band at apical third, and narrow apex 
dorsally and posteriorly brown; middle fe- 
mur with apical half to two-thirds anteriorly 


913 


and apical fourth posteriorly brown; hind 
femur with apical half to two-thirds ante- 
riorly and posteriorly and apical third to 
half dorsally and ventrally brown. Middle 
femur with 2 anterior, 2 anteroventral, and 
1 preapical, dorsoposterior bristles. Hind fe- 
mur with 3 anterior and 5 anteroventral 
bristles, all long and yellowish, most stout; 
anteroventral bristles at middle two-thirds; 
1 posteroventral bristle basally plus 6 bris- 
tles on a raised posterior tubercle at middle 
of hind femur; all posteroventral bristles 
quite thick from base to apex, only slightly 
flattened with round apices, most or all 
brown; HFWLR 1/2.8—1/3.0. Tibiae mostly 
yellow with yellowish bristles laterally and 
brown bristles apically; narrow apices and 
anterior surfaces of all tibiae brown; fore 
tibia with only narrow brown stripe anteri- 
orly. Hind tibia short, two-thirds as long as 
hind femora and four-fifths as long as hind 
tarsus; thick, constricted preapically, apex 
abruptly wide, clublike. Basal tarsomere of 
fore and mid tarsi yellow with narrow apex 
brown; basal tarsomere of hind tarsus yel- 
lowish basally grading to brownish yellow 
apically; apical 4 tarsomeres of all tarsi 
brown; fore and middle tarsi with 5 and 4 
yellow bristles respectively; hind tarsus 
with 2 yellow bristles. 

Abdomen: Yellow brown basally, brown 
apically; mostly yellowish gray tomentum 
and yellowish setae present; terga brown to- 
mentose with brown setae dorsally. 

Terminalia (Figs. 1-5): Epandrium with 
a short, asymmetrical, apical hook; apical 
third with abundant short, bristly setae. 
Ventral lamella with a short, sclerotized, 
preapical, tongue-like process, sides slight- 
ly curved downward; dense, long, yellow 
setae present. Gonostylus with prominent 
wide base and a short, narrow dorsal pro- 
cess. Aedeagus with broad hooded sheath, 
distiphallus strongly arched downward with 
only apex exposed. Gonocoxite with long, 
thin, erect flanges, one along inner margin 
and another more lateral a long, horizontal 
process with pointed apex present. Hypan- 


914 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


2D 


if il 


0.5 mm 


1 0.5 mm D) 


ee as 


3 0.5 mm 


0.3 mm 


SD 


Ommatius mariae, male terminalia. 1, Dorsal view. 2, Lateral view. 3, Ventral view. 4, Gonostylus. 


Figs. 1—S. 
5, Aedeagus, lateral view. 


VOLUME 102, NUMBER 4 


8 0.2 mm 


Figs. 6-9. 
fork. 


drium strongly produced apically, apical 
margin subtruncate, with rounded corners. 

Female.—As male, differing as follows: 
Body 13.6 mm; wing 10.0 mm; FHWR 1/ 
6.9; cell ml WR 1.0:2.1:1.4; HFWLR 1/ 
5.6. Head: Face with 2 brownish yellow to 
brownish bristles. Occiput with 1-2 brown 


915 


9 0.2 mm 


Ommatius mariae, female terminalia. 6, Dorsal view. 7, Ventral view. 8, Spermatheca. 9, Genital 


postocular bristles. Thorax: Tomentum 
largely dull yellowish. Wing: Crossvein r- 
m longer than vein CuAl+Ms3. Base of cell 
r4 at apex of cell d. Legs: Fore coxa with 
several, long, thin bristles. Middle and hind 
femora black on apical half or slightly less 
and with all or most anterior bristles black. 


916 


Hind femur slender, not especially swollen, 
without a mound-like tubercle, and bristles 
acutely pointed; all ventral bristles yellow 
and distributed in well defined rows. Tibiae 
with narrow apex and anterior surface 
black; hind tibia black except yellow on 
basal half anteriorly. Hind tibia slender, 
only gradually wider apically, apex not un- 
usually wide or flat. Tarsi largely or entirely 
black with black bristles; fore and middle 
tarsi with basal tarsomere mostly yellow, 
narrow apex black; only 2—3 yellow bristles 
present. 

Abdomen: Apical margin of tergite 7 
with 2 midlateral black bristles, 1 of these 
contrastingly long and stout. Tergite 8 
shiny, jet black, apical corner somewhat an- 
gular, oblique; a long, midlateral, black 
bristles present posterior to apical margin 
and 2-3 shorter bristles more basally, these 
black or yellowish. Tergite 9 dorsally with 
narrow base sclerotized, less than one-fifth 
as long as cercus, corners contrastingly 
wide, extending around base of ventral la- 
mella forming a wide sclerotize bridge in 
dry specimens. Cercus unusually narrow 
with apex slightly emarginate. Ventral la- 
mella narrowly sclerotized along inner mar- 
gin. Sternites 5—7 with 3—4 long, stout bris- 
tles laterally, mostly yellowish, those in api- 
cal corner of sternite 7 black and contrast- 
ingly long. Sternite 8 shiny jet black with 
a transverse row of yellow setae and bris- 
tles; 4 bristles medially, darker, light amber, 
contrastingly long and thick, extending for- 
ward to subapex of ventral lamella. Sper- 
mathecae obovate; ducts long and thin with 
bases of ducts separate, each originating 
from very short, common duct between 
base of arms of genital fork. Genital fork 
largely membranous, apical third of each 
arm, narrow, slightly sclerotized. 

Types.—Holotype 6, BAHAMAS: 
Eleuthera, Rainbow Bay, 21.x.85, J. R. Wi- 
ley (FSCA). Allotype 2, same data as ho- 
lotype except: 29.vi-6.vii.1990, J. R. & S. 
C. Wiley (FSCA). Paratype: 1 6, same data 
as holotype (FSCA); 1 6, BWI, Nassau, 
3.xii.61, N. A. Roeff (USNM). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Etymology.—Latin mariae, for Mary, in 
memory of a loving, devoted wife and 
mother. 

Remarks.—Ommatius mariae and O. ha- 
nebrinki (holotype and allotype, USNM; 
paratypes, AMNH, TUMZ; others CMNH/ 
FSCA) are very similar species, almost 
identical in general habitatus, and differ 
markedly from other known New World 
species. They are readily separated from 
each other by the uniquely modified struc- 
tures of the terminalia. In addition, the 
whitish ocellar setae and the flat, mostly 
brown bristles with round tips, clustered on 
a raised, posterior tubercle on the hind fem- 
ora further characterize the male. In O. ha- 
nebrinki, these bristles are flatter, yellow, 
and distributed more in a linear row on a 
lower tubercle. The anterior surface of all 
tibiae in the female of O. mariae is mostly 
black whereas the tibiae are yellow with 
narrow dark apices in O. hanebrinki. The 
basal tarsomere of the hind tarsus is black 
in O. mariae whereas it is much lighter, 
brownish yellow in O. hanebrinki. Further- 
more, in O. mariae segment 8 is shiny, jet 
black; tergite 8 has scattered bristly setae 
dorsally and stout bristles laterally, none are 
present along the apical margin. Sternite 8 
medially has a transverse row of numerous, 
short setae and four contrastingly long bris- 
tles. In O. hanebrinki, segment 8 is much 
lighter brown; tergite 8 has mixed bristly 
and abundant, short, recumbent setae dor- 
sally, and several, stout bristles are present 
along the apical margin. The transverse row 
of vestiture on sternite 8 medially consists 
of 8—10 stout bristles and sparse, short se- 
tae. 

Ommatius hanebrinki (1983), O. hispidus 
Scarbrough (1985a), and O. mariae are 
similar in several regards, and differ from 
all known species from Cuba and the Ba- 
hamas. This group, i.e., hanebrinki group, 
is readily recognized by the following com- 
bination of characters of the male: 1) 
Strongly inflated hind femur; 2) a series of 
flat bristles with rounded tips on the pos- 
terior margin of the hind femur; 3) an api- 


VOLUME 102, NUMBER 4 


cally clubbed [hanebrinki and mariae] or 
long digitate process [hispidus] on the hind 
tibia; 4) the unusually long, blade-like pro- 
cesses of the gonocoxite in males; 5) the 
unusually stout bristles on the apical ab- 
dominal segments; and 6) the dorsally 
membranous tergite 9 which wraps below 
the bases of the ventral lamella, and the un- 
usually strongly, apically pointed sternite 8 
with stout bristles posteriorly in the female. 


Ommatius lineolatus Scarbrough 


Ommatius lineolatus Scarbrough 1988: 90— 
94. 5 Holotype, 2 allotype. Type locality 
Cuba (USNM). 


Specimens examined.—BAHAMAS: 1 
2 of O. lineolatus, Bimini (FSCA); 1 6 of 
O. lineolatus, Long Island (FSCA). CUBA: 
Holotype 6 and 2 of O. lineolatus Scar- 
brough (USNM). 

Remarks.—Ommatius lineolatus, a Cu- 
ban species, is reported here from Long Is- 
land in the Bahama Islands and increases 
the number of species of Ommatius to 6 
from this region. The species is readily rec- 
ognized by its small size (S—6 mm), yellow- 
ish brown color of the legs, 6 ventral setae 
distributed the length of the antennal style, 
and combined characters of the terminalia. 
It is otherwise quite similar to O. abana 
(types series, AMNH), and may prove 
eventually to be that species as more ma- 
terial is located and studied. This type series 
of O. abana consist of only 3 females, i.e., 
males are unknown. The species is large (9 
mm), the body is jet black, the legs are am- 
ber or reddish with black markings, and the 
style has only four long setae near the tip. 


Ommatius dignus Scarbrough, 
new name 


Ommatius dimidiatus Scarbrough 1985b: 
647-650. Figs. 9-14 (primary junior 
homonym of Ommatius dimidiatus Mac- 
quart 1850: 394, Asilidae); (types 
GPAC). 


The binomen Ommatius dimidiatus Mac- 
quart (Macquart 1850) was first used for a 


Ov 


species in the Australian region. It is now 
reported from Tasmania and Australia 
(Daniels 1989). Later (Scarbrough 1985b), 
the same binomen was applied to a new 
Neotropical species from Dominica, Lucia, 
and Martinique in the Lesser Antilles. I pro- 
pose dignus (L, meaning worthy or fit) as 
a new replacement name. 


Ommatius fimbrillus Scarbrough, 
new name 


Ommatius fimbriatus Scarbrough and Poin- 
ar 1992: 13-16. Figs 1—4 (primary junior 
homonym of Ommatius fimbriatus Hardy 
1949: 301, Asilidae); (types GPAC). 


Unfortunately, Scarbrough and Poinar 
(1992) were unaware that the binomen Om- 
matius fimbriatus had already been used for 
a species of an asilid fly from Australia 
when they used it for a fossil species in 
Dominican amber. I propose fimbrillus (L, 
meaning a fringe) as a new replacement 
name for it. 


ACKNOWLEDGMENTS 


My sincere thanks to David Grimaldi, 
AMNH, New York; Chen Young, CMNH, 
Pittsburgh; E Christian Thompson, System- 
atic Entomology Laboratory, USDA, Wash- 
ington, DC; and Gary Stark, FSCA, Gaines- 
ville, FL, for the loan of specimens. I also 
thank FE Christian Thompson and Gary 
Steck for their help and cooperation during 
my visit to examine specimens; and to Drs. 
William Grogan, Department of Biological 
Sciences, Salisbury State University, Salis- 
bury, MD, and E Geller-Grimm, Museum 
Wiesbaden (MWNH), Department of Nat- 
ural Sciences, Germany, for their invaluable 
reviews of this manuscript. 


LITERATURE CITED 


Curran, C. H. 1953. The Asilidae and Mydaidae of the 
Bimini Islands, Bahamas, British West Indies (Dip- 
tera). American Museum Novitates 1644: 5—6. 

Daniels, G. 1989. Family Asilidae, pp. 326-349. In 
Evenhuis, N. L., ed., Catalog of the Diptera of the 
Australasian and Oceanian Regions. Bishop Mu- 
seum Press & E. J. Brill, 1,155 pp. 

Hardy, G. H. H. 1949. Miscellanous notes on Austra- 


918 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


lian Diptera. XIV. Venation and other notes. Pro- 
ceedings of the Linnean Society of New South 
Wales 73(1948): 298-303. 

Macquart, P. J. M. 1850. Diptéres exotiques nouveaux 
ou peu connus. 4. Supplément. - Mémoires de la 
Société Royale des Sciences, de |’Agriculture et 
des Artes (1849): 309—479, 14 plates. (365—400, 
plates. 6—9). Lille. 

McAlpine, J. E 1981. Morphology and terminology— 
Adults, pp. 9-63. In McAlpine, J. FE et al., Manual 
of Nearctic Diptera, Vol. 1 Ottawa. Monograph 
No. 28, 674 pp. 

Scarbrough, A. G. 1985a. New Ommatius Wiedemann 
(Diptera: Asilidae) from Cuba and the Bahamas. 
Journal of the New York Entomological Society 
93: 1226-1239. 

. 1985b. Ommatius (Diptera: Asilidae) in the 


lesser Antilles. Proceedings of the Entomological 

Society of Washington 87(3): 641—655. 

. 1988. New robber flies (Diptera: Asilidae) 

from Mayaguana Island, Bahamas, and Cuba. En- 

tomological News 99(2): 90—94. 

. 1997. New and old species of Ommatius Wie- 
demann (Diptera: Asilidae) from Hispaniola. In- 
secta Mundi 11: 9—24. 

Scarbrough, A. G. and R. Rutkauskas. 1983. A new 
species of Ommatius Wiedemann (Diptera: Asili- 
dae) from San Salvador, the Bahamas. Proceed- 
ings of the Entomological Society of Washington 
85: 144-151. 

Scarbrough, A. G. and George O. Poinar. 1992. Upper 
Eocene robber flies of the genus Ommatius (Dip- 
tera: Asilidae) in Dominican Amber Insecta Mun- 
di. 6: 13-18. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 919-923 


A NEW SPECIES OF CHLORONIA BANKS (MEGALOPTERA: 
CORYDALIDAE) FROM SOUTHEASTERN BRAZIL, 
WITH A KEY TO THE SPECIES OF BRAZIL 


ATILANO CONTRERAS-RAMOS 


Corregidora 460-7, Amp. Miguel Hidalgo, 14410 Tlalpan, México, D.F, México 
(e-mail: acontreras_ramos @ hotmail.com) 


Abstract.—The Neotropical dobsonfly genus Chloronia (Megaloptera: Corydalidae: 
Corydalinae) contains 14 previously described species. In this paper, Chloronia pennyi, 
a new species from Minas Gerais, Brazil, is described and illustrated. In general appear- 
ance C. pennyi resembles most the Andean C. bogotana Weele, especially in the small 
fuscous markings on the head and the pair of fuscous bands on the pronotum. In the new 
species, the antennae are entirely yellow (apically infuscate in C. bogotana), the wings 
are mostly pale (patterned in C. bogotana), and the dark bands on the pronotum are 
continuous (nearly meeting in middle in C. bogotana). A key for adult males of the four 
currently recognized Brazilian species is included. New distribution records are given for 
C. corripiens (Walker) in southeastern Brazil. 


Key Words: 
sonfly, taxonomy, key 


The Neotropical genus Chloronia is dis- 
tributed from northwestern and northeastern 
Mexico, southward into western and south- 
eastern South America, including some of 
the Lesser Antilles. Individuals of this ge- 
nus are rather small dobsonflies (forewing 
length 24—50 mm) with a characteristic col- 
or (yellow with black spots), which allows 
for easy recognition. After the revision by 
Penny and Flint (1982), Flint (1991) clari- 
fied the identity of Chloronia bogotana 
Weele and added two new species from 
Costa Rica (Flint 1992), and Contreras-Ra- 
mos (1995) described two new species each 
from Ecuador and Guatemala. In all, 15 
species of Chloronia, including the one de- 
scribed in this paper, are currently recog- 
nized. Larvae of Chloronia have been di- 
agnosed by Penny and Flint (1982) and 
Contreras-Ramos and Harris (1998). Ac- 


Chloronia, new species, Megaloptera, Brazil, Neotropics, Corydalidae, dob- 


counts on habitat of a few species have 
been given by Penny and Flint (1982), Ge- 
ijskes (1984), and Contreras-Ramos (1999). 

In this paper, a new species of Chloronia 
from southeastern Brazil is described and 
illustrated. In 1998, five specimens of the 
new species were collected in two localities 
in Minas Gerais during caddisfly survey 
work by Ralph W. Holzenthal (UMSP) and 
colleagues. General similarity of the distinct 
Brazilian series with the distantly distrib- 
uted C. bogotana (Andes of Bolivia, Co- 
lombia, Ecuador, and Peru). indicated high 
possibilities for the Brazilian specimens to 
belong in a new species. This was corrob- 
orated after careful examination of the male 
genitalia of the Brazilian series. Additional 
distributional records for C. corripiens 
(Walker) in southeastern Brazil, and a key 
for the identification of the four Chloronia 


Rig e 


Habitus of Chloronia pennyi, holotype. 


species currently recorded in Brazil also are 
presented in this paper. 

The genitalic and venational terminology 
used here follows that of Glorioso (1981), 
as modified by Contreras-Ramos (1998). 
Specimens were originally preserved in 
80% ethanol. Three males were spread and 
pinned for dry preservation. Genitalia were 
dissected, cleared, and stored by standard 
methods, as explained by Contreras-Ramos 
(1998). Specimens will be deposited at the 
entomological collections of the Museu de 
Zoologia, Universidad de Sao Paulo, Brazil 
(MZSP); Department of Entomology, Uni- 
versity of Minnesota, St. Paul (UMSP); and 
the National Museum of Natural History, 
Smithsonian Institution, Washington, DC 
(NMNH). 


Chloronia pennyi Contreras-Ramos, 
new species 
(Figs. 1—6) 
Diagnosis.—This species, together with 
Chloronia corripiens (Walker) and C. plau- 
manni Penny and Flint, appears to have a 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


phylogenetically basal position in the ge- 
nus. All three have an unmodified ninth 
sternite, a sparsely setose ninth tergum 
(lacking small clusters of spinous setae pre- 
sent in most species), as well as conspicu- 
ous dorsolateral pregenital sacs between the 
eighth and ninth abdominal segments. 
However, on the basis of general appear- 
ance, the new species is most similar to C. 
bogotana. Both have small posterolateral 
spots on the head (Flint 1991, figs. 5, 6) 
and elongate dark spots on the pronotum. 
In C. bogotana, nevertheless, the pronotal 
spots are discontinuous, not meeting in the 
middle (Flint 1991, figs. 5, 8), whereas in 
the new species they form a pair of contin- 
uous lateral bands (Fig. 6). Forewing col- 
oration in C. bogotana is patterned, with 
most crossveins dark (Flint 1991, figs. 7— 
9). In the new species, forewings are mostly 
clear with few crossveins slightly dark (Fig. 
1). With respect to genitalic characters, the 
10th sternite of C. pennyi resembles that of 
C. convergens Contreras-Ramos. In both, 
10th sternite lobes are sclerotized and su- 


Figs. 2-6. 
eral. 6, Pronotum, dorsal. 


bquadrate, but in the new species lobes lack 
apical spines and the 10th sternite has only 
slightly developed anterolateral projections 
(Fig. 4). 

Description of adult (Figs. 1, 5—6).— 
Head width, ¢ 3.6—4.1 mm (average 3.8 
mm, n = 3), 2 4.1-4.3 mm (average 4.2 
mm, n = 2); forewing length, 6 27.4—29.4 
mm (average 28.3 mm, n = 3), 2 34.3- 


ee . ae 


25 
Senet 


7 


Chloronia pennyi. 2, Male genitalia, dorsal. 3, Same, ventral. 4, 10th sternite. 5, Head, dorsolat- 


34.7 mm (average 34.5 mm, n = 2). Color 
generally pale yellow with fuscous spots 
and bands. Head pale yellow, mandible yel- 
lowish brown with teeth and outer side dark 
reddish-brown. Labrum subquadrate with 
pair of long, flattened setae. Clypeal margin 
nearly straight. Compound eyes and base of 
ocelli dark. Postocular spine blunt, colored 
as head. Pattern of fuscous wide-elongate 


922 


spot at postocular plane, thin spot at adja- 
cent ridge, pair of posterior spots, and 
round spot at occiput, on each side of head 
(Fig. 5). Antenna 37 to 44-segmented, fili- 
form, yellow, with at most last segment in- 
fuscate. Maxilla yellow with 5-segmented 
palp, last two segments pale brown. Labial 
palp 4-segmented, yellow. 

Pronotum yellow, with pair of dark, con- 
tinuous, longitudinal bands (Fig. 6). Meson- 
otum without fuscous spots. Legs yellow, 
tarsal claws brown. Forewing pale yellow, 
hyaline, with 26—27 costal crossveins, 0—2 
forked, a few in proximal % of wing with 
dark ends. Veins mostly yellow, R,-Rs 
crossveins, forking of M, and few basal 
crossveins finely fuscous. Posterior margin 
of wing with subtle grayish maculations. 
Anterior 2A cell variably with a spot to 
only a slight maculation. Hindwing pale 
yellow, hyaline, but 2nd r brown. 

Male genitalia (Figs. 2—4): Ninth tergum 
subtriangular, finely and sparsely setose, 
without patches of spinous setae; V-shaped 
internal inflection reaching midlength of 
tergum. Tenth tergites about 2.5 times as 
long as ninth tergum, subcylindrical, blunt- 
ly tapering, basal 7, divergent, apical /, di- 
rected posteriorly, finely and evenly setose 
(Fig. 2). Ninth gonostylus incurved, fusi- 
form, anteroventral margin slightly more 
convex, with sharp apical point. Ninth ster- 
num moderately sclerotized, subquadrate, 
with well developed posterolateral lobes, 
slightly convex posteromesally (Fig. 3). 
Membrane between 9th and 10th sternites 
eversible, broadly bilobate, thickened. 
Tenth sternite convex, with small, sharply 
pointed anterolateral projections; lobes 
sclerotized, papilliform, subequal in width 
and length, sparsely and conspicuously se- 
tose (Fig. 4). 

Material examined.—Holotype 3: BRA- 
ZIL. Minas Gerais: Serra do Cip6, Rio Cip6 
in Cardeal Mota (Cach. Baixo), 19°20.553’S, 
43°38.531'W, el. 750 m, 10—15.ii.1998, Hol- 
zenthal, Paprocki, Huisman [head width = 
3.6 mm, forewing length = 28.0 mm] 
(MZSP). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Paratypes: Same data as holotype, 1 @ 
(MZSP), 1 do, 1 2 (UMSP); Minas Gerais: 
confluence Rio Peixe & Rio Preto do Itam- 
bé,, 19°17.525'S, 43°15.457'W,, ely S00mm: 
4.11.1998, Holzenthal & Paprocki, 1 ¢d 
(NMNH). 

Etymology.—This new species is gladly 
dedicated to Norman D. Penny from the 
California Academy of Sciences, in recog- 
nition of his extensive contributions to Neo- 
tropical neuropterology, as well as ac- 
knowledging his support for the author’s 
graduate research projects. 


KEY TO KNOWN BRAZILIAN SPECIES OF 
CHLORONIA 
(MODIFIED FROM PENNY AND FLINT 1982) 


1. Head with pair or few small fuscous spots pos- 
teriorly 
— Head unicolorous or only with lateral margin 
INfUSCAIE: 2 sc)y5 a8 Yolo aca ee 2 
2. Lateral margin of head fuscous (Penny and 
Flint, 1982. fig. 44). ates Se 
C. plaumanni Penny and Flint 
— Lateral margin unmarked (Penny and Flint 
OS Dh oe AD) eee gta C. corripiens (Walker) 
3. Pronotum with two anterior and two posterior 
fuscous spots, wings patterned (Penny and 
Flint 1982, fig. 48) .. C. hieroglyphica (Rambur) 
— Pronotum with two fuscous, longitudinal bands 
(Fig. 6), wings mostly clear (Fig. 1) ..... 
bo, Sasa tY ans eg USS C. pennyi, new species 


ADDITIONAL MATERIAL EXAMINED 


Chloronia corripiens (Walker).—BRA- 
ZIL. Minas Gerais: Serra do Cip6, Cardeal 
Mota, Cachoeira Veu da Noiva, 19°18.912'S, 
43°36.260'W, el. 800 m, 12.11.1998, Holzen- 
thal & Paprocki, 1 ¢6 (UMSP); Parana: Rio 
Mae Catira, 10 km N Porto de Cima, 
25°21.821'S,. 48°52.473'W, el. 200iimies— 
9.xii.1997, Holzenthal & Huisman, 3 °& 
(UMSP). 


ACKNOWLEDGMENTS 


Thanks to Ralph W. Holzenthal (UMSP) 
for calling my attention to a series of dob- 
sonfly specimens collected through his cad- 
disfly survey work in Brazil, among which 
the new Chloronia species was found. 
Thanks also to Philip J. Clausen for curat- 


VOLUME 102, NUMBER 4 


ing dobsonflies (UMSP) and processing a 
loan to me. A brief research visit to the 
University of Minnesota was funded by the 
Instituto de Biologia of the Universidad Na- 
cional Aut6noma de México. The hospital- 
ity of Fernando Mufioz-Quesada (UMSP) 
and family is greatly appreciated. Finally, 
thanks to David E. Bowles (Texas Parks 
and Wildlife Department) and an anony- 
mous reviewer for improving the quality of 
the manuscript. 


LITERATURE CITED 


Contreras-Ramos, A. 1995. New species of Chloronia 
from Ecuador and Guatemala, with a key to the 
species in the genus (Megaloptera: Corydalidae). 
Journal of the North American Benthological So- 
ciety 14: 108-114. 

1998. Systematics of the dobsonfly genus 

Corydalus (Megaloptera: Corydalidae). Thomas 

Say Publications in Entomology: Monographs. 

Entomological Society of America, Lanham, 

Maryland, 360 pp. 

1999. Mating behavior of Platyneuromus 

(Megaloptera: Corydalidae), with life 


history 


923) 


notes on dobsonflies from Mexico and Costa Rica. 
Entomological News 110: 125-135. 

Contreras-Ramos, A., and S. C. Harris. 1998. The im- 
mature stages of Platyneuromus (Corydalidae), 
with a key to the genera of larval Megaloptera of 
Mexico. Journal of the North American Bentho- 
logical Society 17: 489-517. 

Flint, O. S., Jr. 1991. On the identity of Chloronia 
bogatana [sic] Weele (Neuropterida: Megaloptera: 
Corydalidae). Proceedings of the Entomological 
Society of Washington 93: 489-494, 

1992. A review of the genus Chloronia in Cos- 
ta Rica, with the description of two new species 
(Neuropterida: Megaloptera: Corydalidae). Pro- 
ceedings of the Biological Society of Washington 
105: 801-809. 

Geijskes, D. C. 1984. Notes on Megaloptera from the 
Guyanas, S. Am., pp. 79-84. In Gepp, J., H. As- 
pock, and H. Holzel, eds., Progress in World’s 
Neuropterology; Proceedings of the Ist Interna- 
tional Symposium on Neuropterology. Graz, Aus- 
tria. 

Glorioso, M. J. 1981. Systematics of the dobsonfly 
subfamily Corydalinae (Megaloptera: Corydali- 
dae). Systematic Entomology 6: 253-290. 

Penny, N. D., and O. S. Flint, Jr. 1982. A revision of 
the genus Chloronia (Neuroptera: Corydalidae). 
Smithsonian Contributions to Zoology 348: 1—27. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 924-928 


A NEW AFROTROPICAL SPECIES OF ALLOGNOSTA OSTEN SACKEN 
(DIPTERA: STRATIOMYIDAE) 


NORMAN E. WOODLEY 


Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- 
ment of Agriculture, % National Museum of Natural History, Smithsonian Institution, 
Washington, DC 20560-0168, U.S.A. (e-mail: nwoodley @sel.barc.usda.gov) 


Abstract.—Allognosta njombe, new species, is described from localities around the 
north end of Lake Nyassa in Tanzania (type locality) and Malawi. A key to the four 
known species of Afrotropical Allognosta is provided. Additional locality records are 
given for A. stuckenbergae Lindner and A. tessmanni Enderlein. 


Key Words: Beridinae, Tanzania, Malawi 


The genus Allognosta Osten Sacken is 
the only genus of the stratiomyid subfamily 
Beridinae in the Afrotropical Region. When 
I reviewed the Afrotropical species (Wood- 
ley 1987) I had available only 12 specimens 
representing three species, although about 
seven additional specimens of A. stucken- 
bergae Lindner were known at that time. A 
small number of additional specimens have 
come to light, including some of a new spe- 
cies, that are reported upon in this paper. 
Since my review of the African species, I 
have published a revision of the world gen- 
era of Beridinae (Woodley 1995) which can 
be consulted for general information and 
cladistic relationships at the generic level 
within the Beridinae as well as an overview 
of the genus Allognosta. 

Terminology and methodology follow 
that used in my previous paper (Woodley 
1987) except that the aedeagal complex is 
now called the phallic complex (Sinclair et 
al. 1994). Specimens studied are from the 
Carnegie Museum of Natural History, Pitts- 
burgh, Pennsylvania, USA (CMNH); Natal 
Museum, Pietermaritzburg, Natal, South 
Africa (NMP); and Naturhistoriska riks- 
museet, Stockholm, Sweden (NRS). The 
sections on material examined under al- 


ready described species treat only speci- 
mens seen since my 1987 paper. 


KEY TO AFROTROPICAL SPECIES OF 
ALLOGNOSTA 


1. Pilosity of eyes long and dense, easily visible 

at low magnifications; occurring only in South 
AtiTiCa wes Sane one A. stuckenbergae Lindner 

— Pilosity of eyes short and sparse, visible only 

at higher magnifications; occurring north of 
South Africa 
. Apical one-third to one-fourth of middle and 
hind femora darkened, strongly contrasting 
with whitish-yellow basal portions; second 
segment of palpus yellow 


Nw 


— Middle and hind femora yellowish, without 
strongly contrasting darker areas; second seg- 
ment of palpus black 

3. Female frons strongly produced, the antennae 
inserted below the prominence; male with me- 
dian lobe of phallic complex similar in diam- 
eter and equal in length to lateral lobes; pleura 
wholly yellowish, only vaguely darker beneath 
wing base A. bwamba Woodley 

— Female frons moderately produced (Fig. 2), the 
antennae inserted at the apex of the promi- 
nence; male with median lobe of phallic com- 
plex in dorsal view (Fig. 5) more slender and 
shorter than lateral lobes; pleura usually 
brownish to brownish-black, nearly concolo- 
rous with scutum, but sometimes paler ..... 
Seas EEN Aare Ae Se eee A. njombe, n. sp. 


VOLUME 102, NUMBER 4 


9, 


y 

\ 

v 

\ 
yh 


i 
1 loner 


> 


Figs. 1-2. 


Allognosta njombe Woodley, new species 
(Figs. 1-7) 


Diagnosis.—This species is in a group 
easily distinguished from A. stuckenbergae 
by having nearly bare eyes. It differs from 
A. tessmanni Enderlein in having all femora 
wholly dark yellow. It is most similar to A. 
bwamba Woodley, but differs in having 
darker pleura, a less strongly produced 
frons in females, and different male geni- 
talia. 

Description.—Male: Head (Fig. 1) 
black, 1.5 time higher than long; eye large, 
upper ommatidia larger than lower ones, 
but size transition not sharply delimited; 
ocellar tubercle moderately prominent; face 
short, receding; frons very small, slightly 
convex; head mostly whitish-gray tomen- 


O25 


Ly Meth. 
wee Dp, 


{ 


SSS 
Nash 


— 


SS 


A= 
\\ \ 
Mines 


/ 


== 
/ 
hg 


LZ 


EZ Wy; 
icp fi fl) yy 
pe 


NO 


Left lateral views of heads of Allognosta njombe. 1, Male. 2, Female. 


tose, sparse on frons, face, and upper oc- 
ciput, almost absent on ocellar tubercle; pi- 
losity sparse, pale on lower occiput and 
gena, more brownish on face; eye with ex- 
tremely short, sparse pilosity, visible only 
at high magnification; antenna 0.8 length of 
head, ratio of segments 4:4:[5:2:3:3:3:3:2.5: 
3.5], scape and pedicel brownish-yellow, 
flagellomeres 1—3 yellow, those beyond 
three brownish; hairs on scape and pedicel 
moderately short, dark; first flagellomere 
with usual subapical dark hairs, other fla- 
gellomeres with a few scattered hairs; pal- 
pus slender, the two segments subequal in 
length, the second slightly clavate; both pal- 
pal segments dark brownish with pilosity 
mostly pale, but with a few dark hairs at 
apex of second segment; proboscis dark 


926 


brownish. Thorax with scutum and scutel- 
lum blackish-brown with slight bronzy re- 
flections, postpronotal lobe slightly paler, 
postalar callus dark yellowish; pleura dark 
brownish, but posterior half of anepister- 
num and katepisternum and entire anepi- 
meron orangish-brown; pleura with pale to- 
mentum but with noticeable bare area on 
ventral half of katepisternum; scutum and 
scutellum entirely, densely pilose with 
short, semi-erect brownish hairs, posterior 
part of scutum and scutellum with some 
longer, more erect hairs intermixed; pleura 
with pale pilosity except anterior half of 
anepisternum and entire meron and katepi- 
meron apilose; coxae dark yellow, anterior 
pair slightly suffused with brown; femora 
dark yellowish, posterior pair a little more 
brownish; tibiae brownish; tarsi dark 
brownish, basal portions of first tarsomeres 
vaguely more yellowish; pilosity on legs 
mostly dark, some pale hairs intermixed on 
femora, entirely pale on coxae, scattered 
longer, erect hairs present on femora and 
tibiae; wing moderately infuscated with 
grayish-brown, pterostigma and veins 
brownish, entire wing set with dense micro- 
trichia; halter dark brownish, basal portion 
of stem paler. Abdomen brownish, tergal 
grooves slightly darker; entire abdomen 
with sparse tomentum; pilosity of tergites 
mostly very short, dark, but longer laterally 
and basally, some hairs on first tergite pale, 
pilosity of sternites pale; terminalia with 
gonocoxites (Fig. 3) moderate in size, mar- 
gins diverging posteriorly; apex of hypan- 
drium moderately produced, the apex of the 
process feebly bilobed; gonostylus with lat- 
eral margin slightly arcuate, otherwise sim- 
ple; phallic complex (Figs. 5—6) arcuate in 
profile, median lobe shorter and more slen- 
der than lateral lobes; epandrium (Fig. 4) 
small, unmodified. Length 5.3 mm. 
Female: Differs from male as follows: 
Head (Fig. 2) 1.2-1.3 times higher than 
long, strongly dichoptic, eye with ommatid- 
ia uniform in size; frons 0.4—0.5 width of 
head at ocellar tubercle, produced at lower 
portion to meet upper facial margin, the an- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


tennae being inserted at anterior-most point 
of head in profile; upper frons with margins 
very slightly diverging ventrally, lower part 
appearing vaguely depressed, overall shiny 
and finely punctate; lower frons very short, 
slightly convex; face short, strongly reced- 
ing; upper occiput margins produced pos- 
terodorsally with sharply rounded margins; 
upper frons and upper occipital margins 
without tomentum, lower frons and face 
with dense, pale grayish tomentum; upper 
frons and occipital margins with moderately 
dense pale golden pilosity; palpus more ro- 
bust than in male, second segment more 
strongly clavate. Thorax with scutum and 
scutellum as in male but with very slight 
metallic purplish reflections; pilosity of scu- 
tum and scutellum shorter than in male, 
uniform in length, entirely pale golden col- 
ored; pleura brownish to brownish-black, 
vaguely paler below wing (uniformly paler 
in specimen from Malawi; see remarks be- 
low); hind coxa sometimes darkened; all 
femora uniformly dark yellowish; tibiae 
brownish-black, approximately basal one- 
third of fore and middle femora yellow, 
hind femur only vaguely pale at extreme 
base; tarsi uniformly brownish-black. Ab- 
domen with tergites with vague purplish re- 
flections; sternites slightly paler in color 
than tergites; furca (Fig. 7) with median ap- 
erture large, emarginate anteriorly; postero- 
medial margin triangularly emarginate me- 
dially; spermathecal ducts unsclerotized 
posteriorly; cerci short, second segment 
about 0.7 length of first. Length 5.6—6.0 
mm. 

Type material.—Holotype ¢ (NRS) from 
Tanzania is labeled: “S TANGANYIKA 
Melando Forest 30 mi S of Njombe 2,450 
m. 10.1962 leg. G. Heinrich(HOLOTY PE 
3 Allognosta njombe N. E. Woodley 
1999.”” The locality of collection is in Ir- 
inga Province in Tanzania. The specimen is 
missing the right front tarsus and the left 
middle tibia and tarsus, otherwise it is in 
excellent condition. The terminalia are 
stored in glycerin in a microvial on the 
specimen pin. Allotype 2, 3 2 paratypes 


VOLUME 102, NUMBER 4 


927 


Figs. 3-7. 
and postgenital segments, dorsal view. 5, Phallic complex, dorsal view. 6, Phallic complex, left lateral view. 7, 
Female furca, ventral view. Abbreviations: ep, epandrium; gc, gonocoxites; gs, gonostylus; hyp, hypandrium; 
llb, lateral lobe of phallic complex; ma, median aperture of female furca; m/b, median lobe of phallic complex; 
pmar, posteromedial margin of female furca. 


(NRS, USNM): Tanzania: Mbeya Province, 
Rungve Mt., 2,600 m, 8—11.1962, G. Hein- 
rich. 3 2 paratypes (NRS): Tanzania: 
Mbeya Province, Rungve Mt., 20 mi. SSE 
Mboya [probably = Mbeya], 2,600 m, 
1962, G. Heinrich. 1 2 paratype (CMNH): 
Malawi: Chitipa District, Jembya Reserve, 
18 km SSE Chisenga, 10-08S, 33-27E, 
1,870 m, 1-10 January 1989, J. Rawlins, S. 
Thompson. 

Distribution.—Known only from high- 


Male and female terminalia of Allognosta njombe. 3, Genital capsule, dorsal view. 4, Epandrium 


land localities in Malawi and Tanzania 
around the north end of Lake Nyassa. 

Etymology.—The species name is a noun 
in apposition based on the name of the type 
locality. 

Remarks.—This species is most similar 
to Allognosta bwamba Woodley. The elon- 
gation of the anterior region of the head in 
females is similar in the two species, al- 
though in A. njombe the antennae are in- 
serted at the apex of the anterior most por- 


928 


tion of the head, not below it. The overall 
structure of the male terminalia is similar in 
the two species, especially the shape of the 
posterior margin of the hypandrium. Al- 
though no phylogenetic analysis has been 
done on Allognosta, it seems probable that 
these two species are closely related. None 
of the other Afrotropical species of Allog- 
nosta have been collected in the vicinity of 
Lake Nyassa. 


Allognosta stuckenbergae Lindner 


Allognosta stuckenbergae Lindner 1961: 1. 


Material examined.—mSOUTH AFRICA: 
1 6, 1 &, Natal, Pietermaritzburg, Town 
Bush, xi.1976, R. Miller (NMP); 1 6, 
Transvaal, 12 km. S Sabi, 2530BB, indig- 
enous bush, 3.xii.1976, R. Miller (NMP); 1 
2, Cape Province, Hogsback, 3226DB, for- 
est and forest margins, 13—16.xu1.1985, J. & 
B. Londt (NMP); 2 2, Cape Province, Tsit- 
sikama Coastal National Park, Stormsriv- 
ermond, 34°02'S, 23°53’E, moist medium 
high coastal forest with Podocarpus, 15— 
19.x.1994, Michael Séderlund, Malaise trap 
(NRS); 1 6, Cape Province, Tsitsikama 
Forest Reserve, 33°58'S, 23°54'E, moist 
high indigenous forest with Podocarpus, 
14—19.x.1994, Michael Sdderlund, Malaise 
trap (NRS); 1 2, Cape Province, Bloukrans 
Pass at Varkrivier, 33°57'S, 23°38’E, coastal 
rainforest in ravine, 14—19.x.1994, Michael 
Séderlund, Malaise trap (NRS). 

Remarks.—All of the new locality data 
fit within the known range of A. stucken- 
bergae in the eastern half of South Africa. 


Allognosta tessmanni Enderlein 


Allognosta tessmanni Enderlein 1921: 182. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Material examined.—UGANDA/ZAIRE: 
12, North of Lake Edward, Gyldenstolpe 
(NRS). 

Remarks.—This specimen was collected 
on an expedition headed by Nils Gylden- 
stolpe along the northern parts of Lake Ed- 
ward from 4 April to 6 May 1921 (Thomas 
Pape, personal communication). 


ACKNOWLEDGMENTS 


I thank Chen Young, Carnegie Museum 
of Natural History, Pittsburgh, Pennsylva- 
nia, USA; David Barraclough, Natal Mu- 
seum, Pietermaritzburg, Natal, South Afri- 
ca; and Thomas Pape, Naturhistoriska riks- 
museet, Stockholm, Sweden, for loaning 
specimens on which this paper is based. 
Thomas Pape also supplied additional in- 
formation on some collection localities. 
Stephen D. Gaimari, Smithsonian Institu- 
tion and Thomas J. Henry and Allen L. No- 
rrbom, both of the Systematic Entomology 
Laboratory, provided comments on the 
manuscript. 


LITERATURE CITED 


Enderlein, G. 1921. Uber die phyletisch alteren Stra- 
tiomyiidensubfamilien (Xylophaginae, Chiromy- 
zinae, Solvinae, Beridinae, und Coenomyiinae). 
Mitteilungen aus dem Zoologischen Museum in 
Berlin 10: 151-214. 

Lindner, E. 1961. Athiopische Stratiomyiiden (Diptera) 
V. Stuttgarter Beitrage zur Naturkunde 68: 1—13. 

Sinclair, B. J., J. M. Cumming, and D. M. Wood. 1994. 
Homology and phylogenetic implications of male 
genitalia in Diptera-lower Brachycera. Entomolo- 
gica Scandinavica 24: 407—432. 

Woodley, N. E. 1987. The Afrotropical Beridinae 
(Diptera: Stratiomyidae). Annals of the Natal Mu- 
seum 28: 119-131. 

. 1995. The genera of Beridinae (Diptera: Stra- 

tiomyidae). Memoirs of the Entomological Soci- 

ety of Washington No. 16, 231 pp. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 929-956 


A NEW GENUS OF THE OROBITIDINAE AND DISCUSSION OF ITS 
RELATIONSHIPS (COLEOPTERA: CURCULIONIDAE) 


Boris A. KOROTYAEV, ALEXANDER S. KONSTANTINOV, AND CHARLES W. O’ BRIEN 


(BAK) Zoological Institute, Russian Academy of Sciences, 199034 St. Petersburg, Rus- 
sia; (ASK) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. 
Department of Agriculture, c/o National Museum of Natural History, Smithsonian Insti- 
tution, Washington, DC 20560-0168, U.S.A. (e-mail: akonstan @sel.barc.usda.gov); 
(CWO) Center for Biological Control, Florida A&M University, Tallahassee, FL 32307- 
4100, U.S.A. 


Abstract.—The new genus Parorobitis of the subfamily Orobitidinae and two new 
species, Parorobitis gibbus (Paraguay: Alto Parana) and Parorobitis minutus (Brazil: 
Curitiba, Parana), are described. The subfamily Orobitidinae is redescribed based on char- 
acters of Orobitis and Parorobitis, and diagnoses of the two genera are provided. Several 
characteristics of the head, thorax, wings and elytra are used to compare the Orobitidinae 
with the supposedly related taxa of Zygopinae, Ceutorhynchinae, and Baridinae. A key 


to these taxa is provided. 


Key Words: 


The subfamily Orobitidinae is comprised 
of two strikingly distinct genera. The Pa- 
learctic Orobitis Germar includes the trans- 
palearctic O. cyaneus Linnaeus which de- 
velops in seed capsules of Viola spp. in 
broad-leaved and mixed forests and O. ni- 
grinus Reitter from the Balkans. A new ge- 
nus of this subfamily is described herein in- 
cluding a new species from Paraguay and 
another from Brazil. 

These new taxa are described and an at- 
tempt is made to place them in the context 
of present-day weevil classification. In the 
last section of the paper, the close affinity 
of the newly described genus with the ge- 
nus Orobitis and preliminary conclusions 
regarding the relationships of the Orobiti- 
dinae are presented. It is clear that exami- 
nation of a much larger sample of taxa and 
characters in combination with rigorous 
analysis may be necessary to propose ro- 
bust hypotheses of relationships. 


Curculionidae, Orobitidinae, new taxa, South America 


As in many other beetle families, termi- 
nology of structures in weevils is fairly in- 
consistent and often is used in a misleading 
way (Thompson 1989). Although the sub- 
ject of this paper is not comparative mor- 
phology or analysis of terminology, it is 
useful to provide a brief summary of the 
sources of terms for the most critical and 
disputed parts of the weevil body. 

To describe metathoracic structures, a 
combination of terms proposed by Matsuda 
(1970) and McHugh et al. (1997) and the 
terms previously adapted for leaf beetles 
(Konstantinov and Vandenberg 1996) is 
used. 

General wing terminology follows Ku- 
kalova-Peck and Lawrence (1993). Some 
unique structures of weevils were named by 
Zherikhin and Gratshev (1995). These 
terms are used for structures which were 
not named by Kukalovd-Peck and 
Lawrence (1993). Terms of internal elytral 


930 


structures are those used by Lyal and King 
(1996). 

Terms proposed by Thompson (1989) are 
used for the spermatheca. The term 
‘“‘pump,”’ widely accepted to describe the 
same part of the spermathecal body at- 
tached to the lobe of the duct and lobe of 
gland (Smith 1979, Konstantinov 1998), is 
used instead of “‘tail.”’ As for the other parts 
of the female genitalia, commonly accepted 
terms are used (Morimoto 1962, O’Brien 
and Askevold 1995, Howden 1995). Mori- 
moto (1962), Lyal (1993), and O’Brien and 
Askevold (1995) are the main sources of 
the names used for different parts of the 
male genitalia. 


Subfamily Orobitidinae 


Description.—Body small, 2—3.5 mm 
long, globose or transversely-subrhomboi- 
dal, prothorax and elytra strongly conjointly 
convex (Figs. 1—4). Black; vestiture dense, 
tightly appressed and inconspicuous or sub- 
erect, composed of white and dark scales. 

Rostrum as (Figs. 3—6, 8) long as or 
slightly longer than prothorax, basally 
weakly curved, apically straight; dorsal 
margin more or less angular in lateral view 
over antennal base. Apical part of rostrum 
in female (Fig. 5) much narrower than basal 
part. Ventral surface of head without pos- 
terior tentorial pits and occipital sutures. 
Subgenal sutures fused (Figs. 10, 11). An- 
tenna inserted at basal third of rostrum 
(Figs. 3, 4, 8, 9). Antennal scrobe oblique, 
ventral margin reaching venter of rostrum 
less than half way between antennal inser- 
tion and eye. Scape stout, short, less than 
half as long as rather long and slender fu- 
nicle with 7 antennomeres (Figs. 16, 17). 
Eye medium-sized, weakly convex. Frons 
as broad as base of rostrum or slightly nar- 
rower, sometimes weakly narrowing in mid- 
dle. 

Mandible (Figs. 12, 13, 15) ventrally 
with 2 visible denticles, third denticle cov- 
ered with apex of rostrum; mandible dor- 
sally separated from maxilla by long ven- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


trolateral appendage of rostrum (Figs. 12, 
15). Labial palpi 2- (Parorobitis) or 3- 
(Orobitis) segmented. Mentum long. 

Prothorax subtrapezoidal with base very 
broadly rounded or nearly straight and 
slightly produced posteriorly for most of its 
length. Sides slightly rounded or nearly 
straight. Basal margins of prothorax and el- 
ytra tightly coupled, sometimes jointly 
raised, but not crenulate. Apical margin of 
pronotum not raised. Postocular lobes well 
developed to absent. Lateral tubercles ab- 
sent. Disc of pronotum flat or moderately 
convex in apical half, without median sul- 
cus. Prosternum excavated, not keeled in 
front of fore coxae. Distance between coxae 
nearly equal to width of rostrum. Mesepi- 
meron not visible from above, flat, deeply 
inserted between base of prothorax and el- 
ytra (Figs. 18, 23). Rostral furrow formed 
by rather high, lamelliform keels behind 
fore coxae on prosternum, sometimes lower 
keels also present on mesosternum. 

Mesoscutellum with tall stalk (Fig. 18). 
Mesosternum between middle and hind 
coxae with broad fold projecting over hind 
coxae. Mesosternum and metasternum 
fused. Internal ridge and border between 
these structures barely visible. Mesendos- 
ternite situated between internal wall of 
coxal cavity and mesosternal ridge. Plate 
nearly as long as wide. Basal and apical ap- 
pendages slender (Fig. 19). 

Scutoscutellar ridges of metanotum not 
connected, directed anteroventrally. Alo- 
cristal ridges not connected in middle (Fig. 
20). 

Metendosternite with moderately narrow 
stalk. Furcal arm slender, apically widening. 
Lateral arm short, situated nearly in middle 
of furcal arm. Anterior and posterior trans- 
verse ventral processes of equal length, 
leaving narrow furrow between (Figs. 21, 
22). 

Elytra slightly longer than broad, with 
strongly convex or obsolete humeral prom- 
inences. Elytra with locking sutural struc- 
ture composed of deep longitudinal groove 


VOLUME 102, NUMBER 4 


Fig. 1. 


Parorobitis gibbus, 3. 


on left elytron (Fig. 18) and longitudinal 
ridge on right elytron (Fig. 27). Basal part 
of suture with two callosities and depres- 
sions between on left elytron. Right elytron 
with two depressions, and with ridge be- 
tween them (Figs. 27, 28, 31, 32). Submar- 
ginal ridge of each elytron also complicated 
(Fig. 29, 30). Apex of elytron with longi- 
tudinal stridulatory file. Preapical groove 
absent (Figs. 35, 36). 

Wings well-developed or slightly re- 


$33) 


duced, with well developed R, radial fis- 
sure, and first radial scelerite (Figs. 25, 26).! 

Legs long. Femora broad from base, un- 
armed, subulate to moderately clavate, shal- 
lowly grooved for reception of tibiae. Tib- 
iae (Figs. 39, 42, 43) in female not unci- 
nate; in male, with short apical mucro (Figs. 


! Zherikhin and Gratshev (1995) state that the Orobi- 
tini are wingless; however, Orobitis cyaneus has short- 
ened but fairly well-developed wings (Fig. 26). 


982 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2—4. 
lateral view. 


2, Orobitis cyaneus, 3, dorsal view. 3, O. cyaneus, 3, lateral view. 4, Parorobitis gibbus, d, 


VOLUME 102, NUMBER 4 


40, 41). Apical comb of setae not or slightly 
extending onto outer margin of tibia. Third 
tarsomere bilobed. Claws separate, moder- 
ately divergent, with long median paired 
contiguous process (Figs. 46—48). 

Abdomen with sterna in one plane. Ster- 
na 1—4 of subequal length, 5 almost as long 
as sterna 3 and 4 together. Sternum | nearly 
completely divided by hind coxae. Hind 
margins of sterna 2—4 parallel, moderately 
curving backwards on sides (Fig. 23). Pleu- 
rite 1 long, separated from rest of tergum 1 
by groove, attached by wide membrane 
with granulated microsculpture to metepis- 
ternum (Figs. 24). Pleura 4—7 combined 
with corresponding tergites. Tergum 8 ex- 
posed in males (Fig. 1). 


Diagnoses of the two genera of the 
subfamily Orobitidinae 


1 (2). Body subspherical, rounded. Humeral 
prominences obsolete. Basal margins of 
prothorax and elytra not raised (Figs. 2, 3). 
Dorsum almost uniformly covered with 
black or bluish black tightly appressed 
scales, shining. Dorsum of rostrum evenly 
curved over antennal base (Figs. 3, 9). 
Frons flat. Pronotum evenly convex, with- 
out median or lateral depressions. Basal 
margin of pronotum almost straight, imper- 
ceptibly notched opposite scutellum. Post- 
ocular lobes absent. Mesoscutellum round- 
ed, as long as broad, weakly and evenly 
convex. Elytra longer than broad, all inter- 
vals flat, striae linear. Lateral margin of el- 
ytron slightly and almost evenly rounded, 
inconspicuously emarginate near base over 
mesepimera. Mesosternum without keels. 


and dark brown spots on pronotum and 
contrasting basal band on elytra, with back- 
ground vestiture composed of dense sub- 
erect to subappressed, linear to lanceolate, 
white scales. Rostral dorsum angular over 
antennal insertion (Figs. 4, 8). Frons mod- 
erately depressed, anterior part sloping to 
base of rostrum and posterior part, to ver- 
tex. Pronotum with two submedian obtuse 
prominences in apical half; laterally un- 
even. Basal margin of pronotum notched in 
middle and bisinuate laterally. Postocular 
lobes from well- to poorly developed. Me- 
soscutellum longer than wide, large, tu- 
berculiform, steeply sloping anteriorly and 
gradually sloping to elytra. Elytra broader 
than long. Odd-numbered intervals weakly 
to moderately convex, costiform near base. 
Striae consisting of medium-sized, deep, 
remote punctures, space between latter un- 
dulate. Intervals very densely, finely punc- 
tate, nearly matt. Lateral margin of elytron 
moderately deeply emarginate over mese- 
pimeron, very shallowly bisinuate behind. 
Mesosternum with well-developed keels 
reaching middle coxae. Fore coxa with ob- 
tuse prominence at apex, separated laterally 
by narrow sulcus. Femora distinctly cla- 
vate, with shallow grooves on ventral side, 
all uniformly covered with scales. Claws 
long, length greater than width of last-tar- 
somere at apex (Figs. 46, 47); median pro- 
cess slightly shorter than claws. Abdominal 
suture 1 with lateral pore under brown 
scaly spot. Pleural area of thorax with 
brown spots. 2.9—3.5 mm.. Neotropical 


Shae tame Carats eam airs Parorobitis, new genus 


Parorobitis Korotyaev, O’Brien, and 


Konstantinov, new genus 


eS) 


2 (1). 


Sides of meso- and metathorax evenly con- 
vex, without prominences or depressions. 
Fore coxa without prominence on inner 
side of apex. Femora nearly parallel-sided, 
with glabrous grooves ventrally. Claw 
short, length less than width of tarsomere 
5 at apex (Fig. 48); median process ca. half 
as long as claw. Abdominal suture | with- 
out lateral pore. Sterna uniformly covered 
with yellowish scales. 2-2.8 mm. Palearc- 
tic. Larvae in seed capsules of Viola spp. 


3 6.6 BMS LCH ORES, ONG. Geer eae oer Orobitis Germar 


Body subrhomboidal, angular humeral 
prominence convex, forming broad, mod- 
erately curved callus (Figs. 1, 4). Basal 
margins of both prothorax and elytra both 
finely conjointly raised. Dorsum with pale 


Type species.—Parorobitis gibbus, new 
species. 

Description.—Small; body short, su- 
brhomboidal, dorsally strongly convex; 
prothorax and elytra conjointly convex. 
Body black, densely clothed with suberect 
to subappressed, elongate scales, dorsally 
with contrasting pattern of white and brown 
scales. Black, funicle and club of antennae 
and apical third of rostrum brown, tarso- 
mere 3 dark brown. 

Head capsule small. Rostrum angularly 
curved at antennal base, ventrally weakly 
arcuate, with dorsum more (in female) or 
less (in male) sharply angular (Figs. 4, 8). 


934 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


* 


ae ' 


Fig. 5—7. Head. 5, Parorobitis gibbus, 3, frontal view. 6, Orobitis cyaneus, ¢, frontal view. 7, P. gibbus, 
view through occipital opening, tentorium and foregut with proventriculus. 


VOLUME 102, NUMBER 4 


10 11 


Fig. 8-11. Head, ¢. 8, 9, Lateral view. 10, 11, Ventral view. 8, 10, Parorobitis gibbus. 9, 11, Orobitis 
cyaneus. 


936 


Basal part of rostrum broader than apical 
(Fig. 5), nearly as broad as fore tibia. Api- 
cal part narrower (Fig. 5); in female, 
straight, cylindrical, shining. Antenna in- 
serted at basal third of rostrum. Scape short, 
stout. Funicle long, antennomeres progres- 
sively shorter apically; club oval, short 
(Figs. 5, 16). Eyes strongly anteriorly di- 
rected. Frons at anterior margin as broad as 
base of rostrum, widened posteriorly, weak- 
ly depressed, slightly, transversely ridged in 
middle. Vertex weakly convex, not carinate. 
Sculpture fine. Tentorium well-developed, 
with moderately broad long stalk attached 
to ventral surface of head and two lateral 
branches attached to anteromedial corner of 
eye (Fig. 7). [It is unclear if these branches 
are homologous with anterior or dorsal 
arms.] Foregut with 8-laminate chitinized 
proventriculus (Fig. 7, 14). [According to 
Crowson (1955) this structure may well be 
plesiomorphic for curculionids. It is also 
known to occur in Cossoninae (Caulophilus 
latinasus Say) (Crowson, 1955), Baridini 
(Pachybaris porosus Lec.) (in the latter spe- 
cies it is much longer and all the laminae 
are lower), and Zygopini (Lobotrachelus 
subfasciatus Motschulsky) (it is generally 
shorter with every lamina being taller pos- 
teriorly and shorter anteriorly). ] 

Left mandible with ventral denticles clos- 
er together than on right mandible. Labial 
palpus 2-segmented. Sides of mentum con- 
vex (Fig. 12). 

Prothorax transverse, trapezoidal. Base 
sinuate, noticeably produced posteriorly 
and enclosed by humeral prominences of 
elytra, shallowly notched anterior to scutel- 
lum and bisinuate laterally. Apical margin 
of pronotum raised. Basal margin raised to 
meet raised basal margin of elytra, finely 
carinate. Hind angles weakly projecting, 
sides weakly convex and strongly converg- 
ing to apical prominences immediately 
above postocular lobes. Apical constriction 
moderately deep on dorsum and gradually 
disappearing towards prominences, which 
slope to anterior margin of prothorax. Sides 
near anterior margin with two angular 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


prominences visible in dorsal view. Punc- 
tures on disc dense, uniform, medium- 
sized, rather deep; intervals between punc- 
tures shining, narrow. Ocular lobe well-de- 
veloped, angular. Fore coxae separated by 
width of rostrum. Prosternum in front of 
coxae not keeled, as long as width of ros- 
trum at base, deeply excavated, with mod- 
erately high lamelliform keels behind cox- 
ae. 

Mesothorax short, transverse (Figs. 18, 
19). Mesosternum with keels reaching an- 
teromedial margin of middle coxae. In lat- 
eral view, mesosternum rather steeply de- 
clivous to metasternum. Scutellum large, 
strongly convex, broadly rounded on top. 
Mesepimeron separated from mesocoxa by 
mesosternal appendage (Fig. 23). Mesepim- 
eron not visible from above, although en- 
tering deeply between bases of prothorax 
and elytron. Distance between mesocoxae 
greater than width of coxa. 

Metathorax much longer than mesotho- 
rax (Fig. 18). Metanotal prescutal mem- 
brane thin and vertical. Alocrista moderate- 
ly narrow, without well-developed lateral 
ridge. Scutum well separated by scutoscu- 
tellar groove forming large “‘pocket”’ lat- 
erally, nearly as large as scutellum. Ven- 
trally, scutoscutellar ridge forming wide 
plate directed anteroventrally and dividing 
metanotal cavity into two almost equal- 
sized compartments (Fig. 20). Scutellar 
groove ending at basal margin of notum. 
Allocristal part of notum containing dorsal- 
ly and ventrally separated cavity with two 
elongate openings. Axillary part of metan- 
otum situated anterolaterad of scutum. Part 
of scutum connected with axillary area 
forming nearly parallel-sided, wide, weakly 
sclerotized appendage. Metasternum form- 
ing thick fold over hind coxa. Hind coxae 
separated from elytra by slightly less than 
width of coxa. Metepisternum convex. 

Elytra much broader than pronotum, 
transverse, strongly narrowing apically, 
with unusually convex humeral prominenc- 
es, separately and rather narrowly rounded 
at apices, and extending over base of py- 


VOLUME 102, NUMBER 4 


Fig. 12-17. 12-14, 16, Parorobitis gibbus, S. 
13, Tip of head, lateral view. 14, Proventriculus. 1 


15, 17, Orobitis cyaneus, 3. 12, Mouth parts, ventral view, 
5, Mouth parts. 16, 17, Antenna. 


938 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


mstr 


msc 


Fig. 18-22. 


Thoracic structures of Parorobitis gibbus. 18, Lateral view with right elytron removed. 19, 


Mesothorax and metasternum (metanotum removed), view of internal structures. 20, Metanotum, ventral view. 
21, 22, Metendosternite (21, dorsal view; 22, ventral view). Abbreviations: alr = allocrista; aly = anterior part 
of longitudinal ventral process; amt = furcal arm; ana = anterior notal appendage; atv = anterior part of 
transverse ventral process; elec = elytral lock; lamt = lateral arm; m = membrane; mesn = mesonotum:; mest 
= stalk of mesoscutellum; msc = mesocoxal cavity; msds = mesendosternite; msm = mesepimeron; msst = 
mesepisternum: mst = mesosternum; mstr = merosternal ridge; mtn = metanotum; mts = 


= = metepisternum; mtt 
posterior part of longitudinal ventral process; pma = postmedial appendage; psm = 


= metasternum; ply = 


VOLUME 102, NUMBER 4 


gidium. Lateral margin of elytron emargin- 
ate at base and very shallowly bisinuate 
posteriorly. Internal surface with four lon- 
gitudinal striae of tiny denticles (Fig. 27). 
Right elytron with locking structure con- 
sisting of narrow ridge and two nearly 
round lateral impressions. Elytral apex cov- 
ered with multidentate median callosities. 
Middle of apex with patch of medially di- 
rected setae (Figs. 27, 31). Striae composed 
of medium-sized, rather deep, sparse, round 
punctures. Intervals 3—4 times as broad as 
striae; odd-numbered intervals weakly to 
strongly convex, densely covered with 
small shallow punctures. 

Wing elongate, with apex slightly darker 
than middle. Humeral field with short C not 
touching Sc. R straight, widening apically. 
Radial fissure indistinct. Radial fold weak. 
First radial sclerite well-developed and 
large, second small. Rr absent. Anal field 
with two weakly developed anal veins (Fig. 
D5): 

Legs long, stout. Trochanters short. Fem- 
ora broad from base, moderately clavate, 
with deep ventral constriction near apex. 
Shallow ventral groove in apical half cov- 
ered with scales as elsewhere. Hind femur 
more swollen in apical part than fore and 
middle femora. Tibiae weakly broadening 
and curved outward apically; fore and mid- 
dle tibiae slightly flattened, hind tibia nearly 
round in cross-section, medially and fron- 
tally compressed apically (Fig. 31). In fe- 
male, tibiae mutic; in male, all armed with 
small curved mucro (Figs. 39—41). 

Apical fringe of setae slightly oblique 
only to outer margin of tibia. Setae dense, 
fine and short. Tarsi of medium proportions, 
tarsomere 3 bilobed, nearly twice as broad 
as 2. Tarsomere 5 moderately widening api- 
cally, slightly more than half length extend- 
ing beyond lobes of tarsomere 3. Claws 


ee 


939 


large, with transversely flattened, longitu- 
dinally curved, apically subacute to acute, 
median process, latter slightly shorter than 
claws (Figs. 46, 47). 

Venter weakly concave in female and 
more strongly so in male. Sterna 1—4 short, 
nearly equal in length. Sternum 5 nearly 
twice as long as sternum 4 (Fig. 23). Small 
indentation between sterna | and 2, slightly 
smaller indentations between other sterna, 
with latter indentations slightly more me- 
dial. Abdominal terga 1—5 short, nearly 
equal in length. Tergum 6 much shorter in 
middle. Tergum 7 slightly longer than terga 
4, 5 and 6 together (Fig. 24). Tergite 8 of 
male with long basal projections (Fig. 57, 
60). 

Male genitalia: Apodemes longer than 
upper part of median lobe. Apical [third? 
fourth] of median lobe bent ventrally, grad- 
ually narrowing apicad. Spiculum gastrale 
long, attached to membrane of tegmen be- 
tween sclerotized parts of sternum 8 (Figs. 
57-62). 

Female genitalia: Tergum 8 elongate, 
weakly sclerotized medially, with moder- 
ately well-sclerotized stripe along border. 
Sternum 8 with two patches of long setae 
apically and short setae between sclerotized 
arms (Fig. 51). Vagina narrow. Coxite and 
stylus narrow, at rest situated inside vagina 
(Figs. 53, 55). Spermatheca with well-de- 
veloped gland and ductal lobes, receptacle 
more than 4X as long as wide (Figs. 52, 
56). 

Etymology.—This masculine generic 
name is the result of combining the generic 
name Orobitis and the Greek prefix para- 
meaning “‘close by, or similar to” and re- 
flects the morphological similarity and 
close relationship of these two genera. 


prescutal membrane; ptv = posterior part of transverse ventral process; scg = scutellar groove; scmt = meta- 
scutellum; sct = metascutum; sctm = mesoscutellum; ssr = scutoscutellar ridge; stmt = furcal stalk; tmt = 


tendons of meso- metafurcal muscles. 


940 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


msst 


<> EEE Ce mts 


26 


Fig. 23-26. 23-25, Parorobitis gibbus. 23, Meso- and metasternum and abdominal sternites. 24, Abdominal 
tergites, pleurites, and metepisternum. 25, Wing. 26, Orobitis cyaneus, wing. Abbreviations: Irs, 2rs = radial 


sclerites; m = membrane; msm = mesepimeron; msst = mesepisternum; p11, 2, and 3 = abdominal pleurites; 
rfi = radial fissure; strm = medial stripe. 


VOLUME 102, NUMBER 4 


Parorobitis gibbus Korotyaev, O’Brien, 
and Konstantinov, new species 
(Figs. 1, 4, 5, 7, 8, 12-14, 16, 18-25, 27, 
29, 31, 35, 39-41, 46, 47, 51-53, 57-59) 


Types.—Holotype d, Paraguay, Puerto P. 
Stroessner, 26—28.XII.1965, Nr. BR 9 (Ma- 
hunka), HMNH. Paratypes. 2 6, 2 2, same 
data as holotype (¢6, 2 HMNH, d USNM, 
2 ZMAS). Paratypes. 6, °, Paraguay: Alto 
Parana, 6 km W. Pto. Pres. Stroessner: 25— 
28.1.1983, leg. E. G. Riley (CWOB) 

Description.—Male: Rostrum nearly as 
long as pronotum, with maximum width at 
antennal insertion, tapering basally and api- 
cally in dorsal view. Dorsal surface with 
sharp median carina disappearing before 
apex (Fig. 5). Lateral carina better devel- 
oped between antennal insertion and head. 
Space between median and lateral carinae 
with row of deep punctures. Weak carinae 
converging on sides above antennal inser- 
tion, but vanishing separately near apex. 
Antenna inserted 0.35 from base of rostrum 
(Fig. 8). Scape shorter than funicular anten- 
nomeres | and 2 together, not reaching eye. 
Funicle rather slender and long, antennom- 
eres progressively becoming shorter, anten- 
nomeres 5 and 6 slightly longer than wide, 
antennomere 7 globose. Club oval, short. 
Setae on flagellomeres moderately long, 
suberect. 

Prothorax 1.65 as broad as long. Pos- 
tocular lobe large, angular. Disc moderately 
convex, with two broad obtuse prominences 
in apical half, separated by shallow longi- 
tudinal depression not extending to basal 
half of disc. Sides of disc with rather deep 
round fovea postero-lateral to apical discal 
prominences, and with shallower fovea be- 
hind anterolateral prominences. Scutellum 
large, 1.4 as long as broad, rounded, very 
strongly convex, with steep anterior slope 
and gentle posterior slope to level of elytral 
suture. Surface of scutellum concealed by 
pale, sand-brown, imbricate scales. 

Elytra 1.2X as broad as long, 1.6% as 
broad as base of prothorax, very strongly 
narrowing apically, with very strongly con- 


94] 


vex rounded humeral prominences. Disc 
strongly convex, more steeply sloping to 
base than to apex. Base of elytra deeply fo- 
veate along basal half of scutellum. Elytral 
striae with medium-sized, deep, sparse 
punctures. Stria 1 at base parallel to antes- 
cutellar margin of elytron and meeting in- 
curved base of stria 2. Striae 3—6 nearly 
straight at base, stria 7 vanishing near the 
posterior margin of swollen interval 7, stria 
8 reaching basal half of humeral promi- 
nence. Intervals 2 and 4 rather strongly 
convex, interval 6 depressed at base, be- 
coming gradually more convex apically. In- 
terval 8 in apical two-thirds slightly more 
convex than others. Preapical prominence 
obtuse and not very convex, stria beneath 
prominence deepened. Intervals matt, 
densely and finely punctate, lacking gran- 
ulations. 

Femora moderately swollen apically. 
Tibiae inconspicuously widening and api- 
cally curved outward. Mucro on middle and 
hind tibiae slightly shorter than tarsal claws; 
on fore tibia, half as long and much finer 
(Figs. 39-41). Protarsomere one about 
1.7X as long as broad, protarsomere two 
1.3X as long as broad, protarsomere three 
0.8X as long as broad, 1.1X as long as 2. 
Tarsomere 5 weakly broadened apically, by 
two-thirds of length extending beyond 
lobes of tarsomere 3. Punctures on femora 
dense, deep, medium-sized, round; on tib- 
iae, more or less elongate. Median claw 
process ca. twice as broad as claw (Figs. 
46, 47). 

Anal sternum flattened in middle one- 
third and covered with suberect to erect 
white hairs, sterna 3 and 4 with long, nar- 
row, curved, suberect white scales along 
hind margin. Pygidium rounded, weakly 
transverse and raised along median line, 
coarsely punctate, and moderately densely 
covered with elongate suberect white 
scales. 

Genitalia (Figs. 57-59): Median lobe 
narrowing apically, abruptly widening hor- 
izontally. Apical third with lateral patches 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 27-28. Internal surface of right elytron. 27, Parorobitis gibbus. 28, Orobitis cyaneus. Abbreviations: 
ag = apical groove of elytron; ape = apical area of elytron; coe = concave area of elyton; f = file; g = 


submarginal ridge; rm = rim; sf = sutural flange. 


VOLUME 102, NUMBER 4 


of setae. Apodeme curved laterally, wid- 
ening before base. 

Head capsule densely covered with par- 
allel-sided or weakly rounded, broad scales; 
vertex with few narrower brown scales in 
middle, and piceous scales on hind margin, 
otherwise with white scales. Basal part of 
rostrum with moderately dense white 
scales, latter half as broad as on vertex, rap- 
idly thinning to antennal insetion; rostrum 
distally with sparse hair-like scales along 
sides to slightly beyond middle. Pronotum 
with moderately dense, narrow-lanceolate, 
brown scales in middle of disc; median de- 
pression with paler scales, often with few 
white scales on bottom. Brown spots on 
discal prominences and near base surround- 
ed by narrow gray lines moderately densely 
covered with hair-like white scales; latter 
also covering sides of prothorax to angular 
line formed by broad white scales, nearly 
concealing pleural area and fore coxa. An- 
terior slope of discal prominences densely 
covered with narrower white scales; apical 
constriction with two gray spots, formed by 
narrow white scales. Sides also with small 
spot of brown scales between hind angles 
and lateral depression. Scutellum dull pale 
brown apically, with white scales at base. 
Base of elytra with band of brown scales; 
intervals 1 and 2, preapical prominences 
and sides in apical half densely covered 
with broad white scales; middle of each el- 
ytron with sparser vestiture of white, linear 
to narrow-lanceolate scales, latter broader 
in striae. Legs mottled with spots of narrow 
and broad white scales, sternum | and base 
of sternum 2 with brown spot on sides. Py- 
gidium with scales not extending from 
punctures, broadly oval to round in basal 
half and elongate, slightly raised near apex; 
brown scales predominant in basal half; 
with white scales, along margins, median 
line, and in apical half. 

Body length 3.1—3.3, width 3 mm. 

Female: Rostrum 1.25 as long as pro- 
thorax, moderately widened and angularly 
curved at antennal insetion, with dorsal out- 
line angular in lateral view; in apical part, 


943 


straight, cylindrical, slender, about *4 width 
of fore tibia. Basal half of rostrum matt, 
densely covered with small shallow punc- 
tures. Median carina well developed, ex- 
tending considerably beyond antennal base; 
lateral carinae finer. Sides of rostrum with 
pair of carinae converging apically from an- 
tennal base to middle of apical part of ros- 
trum. Antennae inserted 0.3 from base of 
rostrum. 

Tibiae without mucro. Protarsomere 1 
about 1.6 as long as broad, protarsomere 2 
nearly as long as broad, protarsomere 3 
twice as long as 2. 

Anal sternum flat. Pygidium slightly lon- 
ger than broad, weakly and rather evenly 
convex, slightly raised along median line in 
apical third, narrowly rounded at apex, 
matt, moderately densely covered with me- 
dium-sized, fairly deep punctures. 

Scutellum dull pale brown. 

Female genitalia: Arms of sternum 8 
moderately long (Fig. 51). Coxite and sty- 
lus slender (Fig. 53), latter nearly cylindri- 
cal. Spermathecal pump and ductal lobe 
forming broad loop. Apex of spermathecal 
pump bent toward ductal lobe (Fig. 52). 

Remarks.—Parorobitis gibbus shares 
most character states with P. minutus. It can 
be separated by the whiter scale pattern of 
the head, larger size, longer arms of ster- 
num 8 (Fig. 51), the slender coxite and sty- 
lus (Fig. 53), the stylus nearly cylindrical, 
the spermathecal pump and ductal lobe 
forming a moderately wide loop, the apex 
of the spermathecal pump bent toward the 
ductal lobe (Fig. 52), the median lobe nar- 
rowing apically with abrupt widening at the 
horizontal part, the apical third with lateral 
patches of setae, and the apodeme widening 
apically (Fig. 57-59). 

Etymology.—This specific epithet is 
based on the Latin adjective gibbus = 
‘*humpbacked, protuberant’”’ and refers to 
the humpbacked appearance and the two 
well-developed pronotal protuberances or 
swellings. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 29-30. 


Parorobitis minutus O’Brien, Korotyaev, 
and Konstantinov, new species 
(Figs. 54-56, 60—62) 


Types.—Holotype ¢, Brazil, Curitiba, 
Parana, I.13.1969, Araucaria forest, leg. C. 
W. & L. B. O’Brien (OZUP). Paratype °, 
same data as holotype (CWOB). 

Description.—Male: Rostrum slightly 
longer than pronotum, with maximum 
width at antennal insertion, tapering basally 
and apically in dorsal view. Dorsal surface 
with sharp median carina disappearing in 
front of apex. Lateral carina better devel- 
oped between antennal insertion and head. 
Space between median and lateral carinae 


Internal surface of right elytron, anterolateral part. 29, Parorobitis gibbus. 30, Orobitis cyaneus. 


without punctures, covered with longitudi- 
nal wrinkles. Lateral carinae above antennal 
insertion parallel to each other, vanishing 
near apex. Antenna inserted 0.37 from base 
of rostrum. Scape shorter than funicular an- 
tennomeres 1 and 2 together, not reaching 
eye. Funicle rather slender and long, anten- 
nomeres progressively becoming shorter, 
antennomeres 5 and 6 slightly longer than 
wide, antennomere 7 globose. Club oval, 
short. Setae on flagellomeres long, suberect. 

Prothorax 1.67X as broad as long. Pos- 
tocular lobe weakly developed. Disc mod- 
erately convex, with single obtuse promi- 
nence in middle. Lateral areas of disc with- 


VOLUME 102, NUMBER 4 


Fig. 31-34. Lock structures of elytron. 31, Parorobitis gibbus. 32, Orobitis cyaneus. 33, Pachybaris porosa. 
34, Lobotrachelus subfasciatus (left elytron). 


out deep fovea, with laterobasal promi- 
nence. 

Scutellum large, 1.67 as long as broad, 
very strongly, roundly convex, with steep 
anterior slope and posterior slope leveling 
to elytral suture. Surface of scutellum con- 
cealed by brownish (in male) or pale sand- 
brown (in female) imbricate scales. 

Elytra together 1.2 as broad as long, 
1.69X as broad as base of prothorax, 
strongly narrowing apically, with very 
strongly convex, rounded humeral promi- 
nences. Disc strongly convex, more steeply 
sloping to base than to apex. Base of elytra 
deeply foveate along basal half of scutel- 
lum. Strial punctures small, moderately 
deep, sparse, nearly entirely covered with 


scales. Stria 1 parallel at base to antescu- 
tellar margin of elytron, meeting incurved 
base of stria 2. Striae 3—6 nearly straight at 
base, stria 7 vanishing in front of preapical 
callosity, stria 8 reaching basal half of hu- 
meral prominence. Intervals 2—6 equally 
convex; intervals 5 and 6 flattening apical- 
ly; interval 8 in apical two-thirds nearly as 
convex as adjacent intervals. Preapical 
prominence convex, stria beneath promi- 
nence deepened. Surface of intervals matt, 
densely and finely punctate, lacking gran- 
ulation. 

Femora moderately swollen apically. 
Tibiae inconspicuously widening and api- 
cally curved outward. Mucro on middle and 
hind tibiae slightly shorter than tarsal claw; 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


\\\ \ \\\\ " 
VAVANAAALL 
\ \\i 


\\ 
\ 


‘ ANN \\\\ ALAA WAAAY At 
\\ ; \A\\\ \ WALA 


ee 


bie 


VOLUME 102, NUMBER 4 


on fore tibia, mucro tiny, scarcely visible. 
Protarsomere one about 2X as long as 
broad, protarsomere two 1.2 as long as 
broad, protarsomere three 0.72 as long as 
broad, 1.3 as long as protarsomere two; 
protarsomere five weakly widened apically, 
extending two-thirds beyond lobes of pro- 
tarsomere 3. Punctures on femora dense, 
deep, medium-sized, round; on tibiae, more 
or less elongate. Tarsal claws with median 
process ca. as broad as claw. 

Anal sternum flattened in middle and 
covered with suberect to erect white hairs, 
ventrites 3 and 4 with long, narrow, curved, 
suberect white scales along hind margin. 
Pygidium rounded, weakly transverse and 
raised along median line, coarsely punctate, 
and moderately densely covered with elon- 
gate suberect white scales. 

Male genitalia (Figs. 57-59): Median 
lobe narrowing apically without abrupt 
widening horizontally. Apical third without 
lateral patches of setae. Apodeme at base 
nearly as broad as at apex. 

Head capsule densely covered with par- 
allel-sided or weakly rounded broad scales; 
vertex with two stripes of narrow brown 
scales in middle, and piceous scales at hind 
margin, otherwise with white scales. Basal 
part of rostrum with moderately dense 
white scales half as broad as those on ver- 
tex, rapidly thinning to antennal insertion; 
rostrum distally with sparse hair-like scales 
along sides to slightly beyond middle. 
Pronotum with moderately dense, narrow- 
lanceolate, brown scales on disc and paler 
scales in middle, with narrow transverse 
stripe of white scales behind middle. An- 
terior slope of discal prominence densely 
covered with wider white scales. Sides also 
with small spot of brown scales above fore 
coxae. Scutellum dull pale brown on top, 
with white scales at base. Outer corner of 
mesepisternum with spot of black scales. 
Base of elytra with band of brown scales, 
intervals 1 and 2, preapical prominences, 
and sides in apical half, densely covered 
with broad white scales, middle of each el- 
ytron with sparser vestiture of white, linear 


947 


to narrow-lanceolate scales, broader in stri- 
ae. Legs mottled with spots of narrow and 
broad white scales, sternum 1 and base of 
sternum 2 with brown spot on sides. Pygid- 
ium with scales not extending beyond punc- 
tures, broadly oval to round on basal half 
and elongate, slightly raised near apex; 
brown scales predominant on basal half, 
white scales along margins, median line, 
and on apical half. 

Body length 2.9—3.0, width 2.9 mm. 

Female: Rostrum 1.25 as long as pro- 
thorax, moderately widened and angularly 
curved at antennal insertion, with dorsal 
outline angular in lateral view; apically 
straight, cylindrical, slender, about *%4 width 
of fore tibia. Median carina shorter than in 
male; lateral carinae well developed. An- 
tenna inserted 0.3 from base of rostrum. 

Tibiae without mucro. Protarsomere one 
about 2.2 as long as broad, protarsomere 
two 1.2X as long as broad, protarsomere 
three 1.17X as long as protarsomere two. 

Female genitalia: Arms of sternum 8 
moderately short (Fig. 54). Coxite and sty- 
lus robust (Fig. 55). Stylus tapering apical- 
ly. Spermathecal pump and ductal lobe 
forming moderately narrow loop. Apex of 
spermathecal pump bent in direction op- 
posite of ductal lobe (Fig. 56). 

Remarks.—Parorobitis minutus shares 
majority of the character states exhibited by 
P. gibbus. It can be distinguished by the 
darker scale pattern of the head, smaller 
size, shorter arms of sternum 8 (Fig. 54), 
robust coxite and stylus (Fig. 55), stylus ta- 
pering apically, spermathecal pump and 
ductal lobe forming rather narrow loop, 
apex of spermathecal pump bent in direc- 
tion opposite of ductal lobe (Fig. 56), me- 
dian lobe narrowing apically without abrupt 
widening horizontally, apical third without 
lateral patches of setae, and apodeme at 
base nearly as narrow as at apex (Fig. 60— 
62). 

Etymology.—This specific epithet is 
based on the Latin adjective minutus = 
‘“‘small’’ and refers to the relatively small 
size of the body. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ali 


Fig. 37-38. Elytral apex with stridulatory file. 37, Pachybaris porosa. 38, Lobotrachelus subfasciatus. Ab- 
breviations: ag = apical groove; pag = preapical groove. 


VOLUME 102, NUMBER 4 949 


Fig. 39-43. Legs. 39-41, Parorobitis gibbus. 42, 43, Orobitis cyaneus. 39, Right metatibia, dorsal view. 
40, 41, Tibial apex. 42, Right metatibia, dorsal view, 43, Right metatibia, ventral view. 


950 


DISCUSSION 


In spite of the sharp differences in ap- 
pearance, Orobitis Germar and Parorobitis 
share many important characters including 
the following: unusual, extremely convex 
shape of body; rostrum weakly curved ba- 
sally, apically straight; ventral surface of 
head without posterior tentorial pits and oc- 
cipital sutures; antennal scrobe oblique, 
with ventral margin reaching venter of ros- 
trum less than half way between antennal 
insertion and eye; mandibles with two den- 
ticles ventrally; labial palpi 2-segmented; 
prothorax subtrapezoidal, with base ex- 
tremely wide and broadly rounded; mese- 
pimera not visible from above, flat, deeply 
inserted between base of prothorax and el- 
ytra; prosternal sulcus formed by rather 
high lamelliform keels behind fore coxae; 
mesoscutellum with tall stalk; mesosternum 
and metasternum fused; elytra with strongly 
convex humeral prominences and sutural 
locking mechanism; apex of elytron with 
longitudinal stridulatory file (Figs. 35, 36); 
wings with well-developed R and radial fis- 
sure; legs long; claws with well developed 
median process (Figs. 46—48); abdominal 
sternum 1 nearly completely divided by 
hind coxae; pleurum 1 long, separated from 
tergum 1 by groove, attached to metepis- 
ternum by wide membrane with granulated 
microsculpture (Figs. 24). One of the most 
convincing pieces of evidence for the close 
relationships between Orobitis and Paro- 
robitis is the structure of the stridulatory 
device. According to Lyal and King (1996), 
Orobitis has a unique file isolated from sur- 
rounding sculpture and placed on a differ- 
ent part of the elytron (Figs. 28, 36). Based 
on these observations, Lyal and King 
(1996) suspected that the file of Orobitis is 
non-homologous with files of other weevils 
(Figs. 37, 38). Parorobitis shares all the 
distinctive characters of the stridulatory file 
of Orobitis (Figs. 27, 35). 

The differentiation between the two gen- 
era 1S most evident in the gestalt. The 
strongly developed relief of the exoskeleton 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


in Parorobitis readily distinguishes it from 
species of Orobitis. Perhaps this indicates 
greater development of thoracic muscles, 
possibly due to a much more active flight 
behavior in the species of the new genus. 

The systematic relationships of the Orob- 
itidinae are uncertain. Colonnelli (1984: 
207) placed Orobitis in the Ithyporinae 
[previously considered a tribe of the Cryp- 
torhynchinae (Hustache 1936)], and cited 
Hustache (1936) as justification for the 
placement. This is actually incorrect as 
Hustache does not deal with Orobitis in the 
Coleopterorum Catalogus, but only cites the 
original placement of Cleogonus nuculus 
(Germar) in the genus Orobitis, which has 
nothing to do with the systematic position 
of Orobitis. Zherikhin and Egorov (1990: 
113) treat Orobitini as a tribe of the Bari- 
dinae (which according to them also in- 
cludes Ceutorhynchini, Zygopini, and Tri- 
gonocolini). 

Zherikhin and Gratshev (1995) elevated 
Baridinae to family, but there are some 
problems with this taxonomic decision. 
Among the synapomorphies of Barididae 
sensu Zherikhin et Gratshev, the presence 
of the median keel on the inner surface of 
metasternum seems the primary character. 
However, this is shared by other apparently 
very different groups and its distribution in 
combination with other characters should 
be studied. The first character in the wing 
structure listed by Zherikhin and Gratshev 
(1995: 773), among the most important 
synapomorphies of the Barididae, is the 
strongly curved basal wing margin. How- 
ever this character state is more developed 
in the Zygopinae and Ceutorhynchinae 
(Zherikhin and Gratshev 1995, Figs. 108— 
121), than in Baridinae (see Figs. 122—132 
in the same work). Trigonocolinae do not 
possess a critical diagnostic character which 
distinguishes Ceutorhynchinae, Zygopinae, 
and Baridinae from the rest of Curculioni- 
dae, i.e., the dorsally visible mesepimera, 
and it is not clear that this character state 
should be considered to be a secondary loss 
in Trigonocolinae as presumed by Zheri- 


VOLUME 102, NUMBER 4 


Fig. 44-50. Legs. 44, Pachybaris porosa, right metatibia, ventral view. 45, Lobotrachelus subfasciatus, right 
metatibia, ventral view. 46—S0. Claw. 46, 47, Parorobitis gibbus. 48, Orobitis cyaneus. 49, Pachybaris porosa. 
50, Lobotrachelus subfasciatus. 


952 


khin and Egorov. The large to very large 
scutellum, the presence of the wax powder 
secretion, and the often carinate elytral in- 
tervals of Trigonocolus do not fit the di- 
agnosis of the Barididae. Also Zherikhin 
and Gratshev (1995) give no evidence from 
the wing structure of close affinity of Tri- 
gonocolinae with the other 4 subfamilies. 
We prefer to consider Trigonocolinae as 
short-bodied representatives of the phyletic 
branch also including Mecyslobini, Mag- 
dalini, and Carciliini (i.e., Molytinae in re- 
cent classifications) rather than to place 
them close to Orobitidinae, Ceutorhynchi- 
nae, Baridinae, or Zygopinae. Each of the 
last four groups can be clearly identified 
and is very different from each other. Ex- 
cept for the Orobitidinae, the other taxa 
have very large numbers of genera and spe- 
cies and are distributed worldwide. Thus it 
seems that additional data is needed for 
treating Baridinae, Ceutorhynchinae, Zyg- 
opinae and Orobitidinae as members of a 
single natural group. Until the presence of 
a longitudinal keel on the metasternum is 
shown to be a true synapomorphy, through 
rigorous character analysis, there is no rea- 
son to consider Barididae sensu Zherikhin 
and Egorov (1990) and Zherikhin and 
Gratshev (1995) to be a natural group. 
Detailed comparative analysis of the taxa 
considered is far beyond the aim of this pa- 
per and needs examination of much greater 
number of taxa and structures. However, we 
would like to summarize here some char- 
acters of the Orobitidinae and their sup- 
posed relatives. We believe that revealing 
clear distinctions will allow better under- 
standing of the groups, rather than lumping 
them together based on characters whose 
synapomorphic value has not been proven. 


KEY FOR DIFFERENTIATION OF THE 
OROBITIDINAE AND WEEVIL SUBFAMILIES 
WITH DORSALLY VISIBLE MESEPIMERA: 


1 (2). Mesepimera not clearly visible from above. 
Meso- and metasterna fused. Abdominal 
sternum | not longer than 2, nearly com- 
pletely divided by hind coxae. Rostrum 


Pal): 


3 (4). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


more or less bent at antennal insertion, 
straight or nearly straight in apical part, al- 
ways narrower than basal part; not separat- 
ed from frons by sulcus. Claws with ap- 
pendages fused into entire flat process half 
as long or nearly as long as claws them- 
selves. Body globose or subrhomboidal, 
very strongly convex dorsally. 1.8—3.5 mm. 
Palearctic and Neotropics ..... 
Mesepimera usually clearly visible from 
above (not visible in Palearctic species of 
Baris Germar developing on crucifers and 
in some southern Asian and South African 
Ceutorhynchinae with subconical, strongly 
apically narrowing prothorax). Meso- and 
metasterna not fused, separated by distinct 
suture. Abdominal sternum | longer than 
2, never nearly completely divided by hind 
coxae. Rostrum not conspicuously bent at 
antennal insertion. Claws simple, toothed, 
or appendiculate, in latter case appendages 
not fused in an entire flat median process. 
Body shape variables aa) see 
Prosternum with median sulcus for recep- 
tion of rostrum, often prolonged on meso- 
or metasternum, and with more or less de- 
veloped keels in front of fore coxae. Males 
always without horn-like projections on 
prosternum before coxae and without fovea 
between them. Head capsule not spherical, 
but transverse or slightly flattened dorso- 
ventrally. Rostrum not separated from head 
capsule by sulcus, neither conspicuously ta- 
pering apically nor dilated to base. Anten- 
nal funicle often with fewer than 7 flagel- 
lomeres. Eyes usually more or less convex, 
always separated on frons (usually by not 
less than twice width of antennal scape). 
Pronotum often with raised anterior margin 
and with lateral (and often also discal) tu- 
bercles. Basal margin of pronotum often 
raised conjointly with basal margins of el- 
ytra and crenulate. Scutellum small to mi- 
nute, never transverse or subcordate. Elytra 
usually broad, slightly longer than broad, 
in tropical species often broader than long. 
Lateral margin of elytron more or less an- 
gularly emarginate above anterior part of 
metepisterna, latter projecting dorsally. 
Many species apterous in temperate zones. 
Hind femora broadest, often saltatorial. 
Tibiae mutic (except in some Zacladus 
Reitter and Scleropterus Schoenherr), mu- 
cro usually developed on middle and hind 
tibiae in males, but often also on fore tib- 
iae; in xerophilous species with narrow tar- 
si, females may have longer mucro. Claws 
usually toothed or appendiculate, often 


Orobitidinae 


955 


VOLUME 102, NUMBER 4 


Fig. 51-56. Female genitalia. 51-53, Parorobitis gibbus. 54—56, Parorobitis minutus. 51, Sternite and tergite 
= sternite 8; 


8. 52, 56, Spermatheca. 53, 55, Vagina with coxites and styli. 54, Sternite 8. Abbreviations: 8 st 
arm of sternite 8; apd = apodeme of sternite 8; cx = coxite; d = duct: dlb = ductal 


gland lobe; sty = stylus; vg = vagina. 


8 t = tergite 8; arm 
lobe; g = gland; glb 


954 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


mdl 


Fig. 57-62. Male genitalia and tergite 8. 57-59, Parorobitis gibbus. 60—62, Parorobitis minutus. 57, 60, 
Tergite 8 and aedeagus, ventral view. 58, 61, Dorsal view. 59, 62, Lateral view. Abbreviations: 8 st = sternite 
8; 8 t = tergite 8; ap = apodeme; mdl = median lobe; spg = spiculum gastrale; tg = tegmen. 


VOLUME 102, NUMBER 4 


4 (3). 


5 (6). 


connate; rarely single. Pygidium exposed 
except in Palearctic Trichosirocalus Colon- 
nelli. Host plants mostly herbaceous or li- 
anas, larvae develop in non lignified tissue. 
1.5—7.0 mm. World-wide including Arctic 
(but not Subantarctic) tundra 


Be es CATS TEM ote Ceutorhynchinae 


Prosternum usually without median sulcus 
for reception of rostrum, without keels in 
front of fore coxae (except in Lobotrach- 
elini, Zygopinae). Prosternum in males 
sometimes with horn-like projections be- 
fore coxae and deeply foveate between 
them. Head capsule spherical or transverse. 
Rostrum usually separated from head cap- 
sule by sulcus and/or more or less strongly 
widening to base, often also tapering and 
flattened apically. Antennal funicle usually 
with 7 flagellomeres. Eyes often flat, broad- 
ly separated or contiguous. Anterior margin 
of pronotum never raised, sides without tu- 
bercles, disc more or less evenly convex, 
sometimes with median tubercle, but not 
sulcate. Basal margins of pronotum and el- 
ytra neither raised conjointly nor crenulate. 
Scutellum small to large, often convex, 
transverse or subcordate. Elytra often much 
longer than broad. Lateral margin of ely- 
tron shallowly sinuate, not conspicuously 
emarginate above anterior part of metepis- 
terna. Wings usually functional except in 
herpetobiont species. Fore femur usually 
broadest, often strongly enlarged and den- 
tate- then fore tibia strongly bent; legs not 
saltatorial. Tibiae uncinate; in males, often 
also mucronate. Claws usually simple, free 
or connate; rarely single. Pygidium ex- 
posed or concealed. Host plants herbs, very 
often lianas and trees, larvae often in lig- 
nified tissue. 0.9-35 mm. Worldwide ex- 
cept northern taiga and tundra ........ 
Eyes contiguous or subcontiguous, very 
large, flat, limited to dorsal half on head 
capsule, not visible ventrally. Head capsule 
at least slightly transverse. Rostrum not 
separated from head capsule by sulcus, 
more or less widening to base, often taper- 
ing and flattened apically. (In Lobotrach- 
elini, Zygopinae, prosternum deeply sul- 
cate, body polished, globose, legs subu- 
late.) Elytra elongate to transverse, usually 
flattened dorsally, never glabrous or metal- 
lic, usually with scales. Wings usually 
functional. Venter more or less strongly 
oblique to apex in lateral view. Femora of- 
ten dentate, fore femur often strongly en- 
larged, then tarsus elongated, especially in 


males. Claws usually free, simple or 


toothed. Predominantly tropical and sub- 
tropical #2. hs. c) Reena are eae ee Zy gopinae 
6 (5). Eyes widely separated on frons, always ap- 
proximate ventrally. Head capsule spheri- 
cal. Rostrum usually separated from frons 
by sulcus. Prosternum rarely deeply sulcate 
for reception of rostrum. Elytra usually 
elongate, in many tropical forms short or 
subglobose, body then often glabrous or 
metallic. Vestiture often reduced or absent. 
Wings reduced in herpetobiont forms. Ven- 
ter not conspicuously oblique apically in 
lateral view. Femora usually unarmed, fore 
femur rarely strongly enlarged. Claws often 
connate, rarely toothed or single. World- 
wide except northern taiga and tundra ... 
spate ackah Fats MEPeteh aes Stee OTE Ac Baridinae 


ACKNOWLEDGMENTS 


We thank O. Merkl of the Hungarian 
Museum of Natural History (HMNH) for 
the opportunity to study material in his 
care. The other depositories of the type ma- 
terial are abbreviated as follows: C. W. 
O’Brien collection, Florida A & M Univer- 
sity, Tallahassee, FL (CWOB); National 
Museum of Natural History, Smithsonian 
Institution, Washington, DC (USNM); Zoo- 
logical Institute, Russian Academy of Sci- 
ences, St. Petersburg (ZIN); Federal Uni- 
versity of Parana, Curitiba, Brazil (OZUP). 
We are also grateful to T. L. Erwin (De- 
partment of Entomology, National Museum 
of Natural History, Washington, DC), A. 
Norrbom, and N. Woodley (Systematic En- 
tomology Laboratory, ARS, USDA, Wash- 
ington, DC) for reviewing this manuscript 
and providing valuable suggestions. Re- 
search by B. Korotyaev was funded in part 
by Russian Federation for Basic Research, 
Grant # 98-04-49763. Research by C. W. 
O’Brien was funded in part by the USDA's 
Cooperative State Research, Education and 
Extension Service (FLAX 910001). 


LITERATURE CITED 


Colonnelli, E. 1984. Notes sur quelques Ceutorhyn- 
chinae de |’ Afrique tropicale (Coleoptera, Curcu- 
lionidae). Annales Historico-Naturales Musei Na- 
tionalis Hungarici 76: 207-238. 

Crowson, R. A. 1955. The Natural Classification of the 
Families of Coleoptera. Lloyd, London, viii+ 187 


956 


pp. [Reprinted from Entomologist’s Monthly 
Magazine, 1950-1954]. 

Howden, A. T. 1995. Structures related to oviposition 
in Curculionoidea, pp. 53—100. Jn Anderson, R. S. 
and C. H. Lyal, eds., Biology and Phylogeny of 
Curculionoidea: Proceedings of a symposium con- 
vened at the XVIII International Congress of En- 
tomology, Vancouver, Canada, July 3-9, 1988. 
Memoirs of the Entomological Society of Wash- 
ington. 14, 174 pp. 

Hustache, A. 1936. Curculionidae: Cryptorrhynchinae. 
Coleopterorum Catalogus. W. Junk, ‘s-Gravenha- 
celle SilWspp: 

Konstantinov, A. S. 1998. Revision of the Palearctic 
species of Aphthona Chevrolat and cladistic clas- 
sification of the Aphthonini (Coleoptera: Chryso- 
melidae: Alticinae). Memoirs on Entomology, In- 
ternational, Associated Publishers, Gainesville. 
429 pp. + 608 figs. 

Konstantinov, A. and N. Vandenberg. 1996. Handbook 
of Palearctic flea beetles (Coleoptera: Chrysome- 
lidae: Alticinae). Contributions on Entomology, 
International 1(3): 237—439. 

Kukalova-Peck, J. and J. EF Lawrence. 1993. Evolution 
of the hind wing in Coleoptera. The Canadian En- 
tomologist 125: 181—258. 

Lyal, C. H. C. 1993. Cryptorhynchinae (Insecta: Co- 
leoptera: Curculionidae). Fauna of New Zealand. 
Manaaki Whenua Press, Lincoln, Canterbury, 
New Zealand. 29, 305 pp. 

Lyal, C. H. C. and T. King. 1996. Elytro-tergal strid- 
ulation in weevils (Insecta: Coleoptera: Curculion- 
oidea). Journal of Natural History 30: 703-773. 

McHugh, J. V., C. J. Marshall, and E L. Fawcett. 1997. 
A study of morphology in Megalodacne heros 
(Say) (Coleoptera: Erotylidae). Transactions of the 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


American Entomological Society 123(4): 167— 
223) 

Matsuda, R. 1970. Morphology and evolution of the 
insect thorax. Memoirs of the Entomological So- 
ciety of Canada 76, 431 pp. 

Morimoto, K. 1962. Comparative morphology and 
phylogeny of the superfamily Curculionoidea in 
Japan. (Comparative morphology, phylogeny and 
systematics of the superfamily Curculionoidea of 
Japan I). Journal of the Faculty of Agriculture, 
Kyushu University 11: 331—373. 

O’Brien, C. W. and I. S. Askevold. 1995. Systematics 
and evolution of weevils of the genus Bagous 
Germar (Coleoptera: Curculionoidea), V. Taxo- 
nomic treatment of the species of the Indian sub- 
continent. Contributions of the American Ento- 
mological Institute 28(5), 182 pp. 

Smith, E. H. 1979. Techniques for the dissection and 
mounting of the male (aedeagus) and female 
(spermatheca) genitalia of the Chrysomelidae 
(Coleoptera). Coleopterists Bulletin 33(1): 93-— 
103. 

Thompson, R. T. 1989. Terminology of the spermathe- 
ca—A protest. Curculio 26: 3—4. 

Zherikhin, V. V. and A. B. Egorov. 1990. Weevils (Co- 
leoptera, Curculionidae) of the Far East of the 
USSR (a review of the subfamilies with a descrip- 
tion of a new taxa). Academy of Sciences, Far 
Eastern Branch, Vladivostok, 164 pp. 

Zherikhin, V. V. and V. G. Gratshev. 1995. A compar- 
ative study of the hind wing venation of the su- 
perfamily Curculionoidea, with phylogenetic im- 
plications, pp. 633-777. In Pakaluk, J. and S. A. 
Slipinski, eds., Biology, phylogeny, and classifi- 
cation of Coleoptera: Papers celebrating the 80th 
birthday of Roy A. Crowson. Muzeum I Institut 
Zoologii PAN, Warszawa. 1,092 pp. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 957-963 


FIRST DOCUMENTED RECORD OF MONOMACHIDAE 
(HYMENOPTERA: PROCTOTRUPOIDEA) IN NEW GUINEA, 
AND DESCRIPTION OF TWO NEW SPECIES 


LUCIANA MUSETTI AND NORMAN FE JOHNSON 


Department of Entomology, The Ohio State University, 1315 Kinnear Road, Columbus, 


OH 43212-1192 U.S.A. 


Abstract.—The occurrence of the genus Monomachus Klug (Hymenoptera: Proctotru- 
poidea, Monomachidae) in New Guinea is documented for the first time. Two new species, 
M. cracens and M. comptus, are described. These are distinguished from the three known 
Australian Monomachus as well as the numerous New World species. 


Key Words: 
asitoid 


The family Monomachidae is a small 
group of parasitic Hymenoptera (Procto- 
trupoidea) with austral disjunct distribu- 
tion. Only two genera have been recog- 
nized, Monomachus Klug and Tetraconus 
Szépligeti. The latter is known only from 
a single female specimen from Brazil. We 
are studying the family with the goal of 
elucidating the phylogenetic relationships 
among species and understanding their 
biogeography. Nineteen species-group taxa 
have been described from the New World 
from Mexico to Chile and Argentina, and 
these were last revised by Schulz (1911). 
Three additional species of Monomachus 
were recognized in the recent revision of 
the Australian fauna (Naumann 1985). 
Masner (1993) noted that the family is also 
to be found in New Guinea, but no species 
have been described. The Papuan material 
in fact represents two species distinct from 
those known in Australia. We describe 
them here because they do not appear to 
be closely related to the numerous Neo- 
tropical species; these will be the focus of 
a separate paper. 


Monomachus, New Guinea, Australia, Proctotrupoidea, Hymenoptera, par- 


MATERIALS AND METHODS 


Specimens for this study from New 
Guinea are found in the collections of the 
American Entomological Institute, Gaines- 
ville, FL (AEIC) and the Bishop Museum, 
Honolulu, HI (BPBM). Australian material 
is found in the Australian National Insect 
Collection, Canberra; Canadian National 
Collection of Insects, Ottawa; the Museum 
of Comparative Zoology, Cambridge, MA; 
The Natural History Museum, London; and 
The Ohio State University, Columbus. 

The mandibles in Monomachidae are re- 
markably diverse in shape. We use the fol- 
lowing terms to describe their structure. 
The mandible is generally divided into two 
areas, a basignath and distignath, separated 
by a subbasal groove. The groove allows 
for greater range of abduction of the man- 
dibles. The distal margin of the groove is 
sometimes sharply marked; this corre- 
sponds to the mandibular fold of Naumann 
(1985) and the Basalfalte of Schulz (1911). 
The lowermost part of the subbasal groove 
is sometimes clearly visible as a sharply de- 


958 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 


fined diagonal sulcus. The basignath is con- 
vex and invaginated into the oral cavity 
when the mandibles are widely opened. The 
nomenclature for thoracic sulci follows 
Huber and Sharkey (1993). 

The label data for all specimens is avail- 
able in electronic format in Hymenoptera 
On-Line at the URL http://ris.biosci.ohio- 
state.edu/hymenoptera (Johnson and Muset- 
ti, in press). The OSUC numbers quoted are 
the individual unique identifiers for the 
specimens in this study. 


Monomachus cracens Musetti and 
Johnson, new species 
(Fig. 1) 


Female.—Measurements for the two fe- 
males are reported in pairs separated by a 
slash; first value refers to female bearing ID 
number OSUC 117646 (holotype @), sec- 
ond value refers to OSUC 117653 (paratype 
2). Fore wing length 5.0/5.8 mm. Color: 
Head, lateral margins of pronotum, central 
disc of mesoscutum on either side of notau- 
li, metanotum, upper portion of mesopleu- 
ron, all of metapleuron, lateral and posterior 
portions of propodeum, entire metasoma 
brown to dark brown; mesosoma otherwise 


Monomachus cracens, \ateral habitus. Scale line = 1 mm. 


light brownish yellow; coxae and trochan- 
ters brownish yellow, legs otherwise dis- 
tinctly darker brown; wing membrane clear. 

Head: Clearly transverse in frontal view, 
width across eyes 2.10/1.96X greater than 
length, head width across the compound 
eyes (2.48 mm/2.69 mm) not differing sig- 
nificantly from head width across gena 
(2.40 mm/2.70 mm respectively). Eye 
height 3.54/4.71 xX malar length. Vertex 
posteriorly with moderately dense deep se- 
tigerous punctures, each separated by ap- 
proximately diameter of one puncture; 
punctures sparser, more irregularly sized 
and spaced near ocelli; punctures densely 
packed, but irregularly shaped near anten- 
nal insertions; vertex otherwise smooth, 
without microsculpture; outer margins of 
ocelli bordered by distinct furrows; vertex 
immediately behind ocelli largely glabrous. 
Occipital carina widely separated from oral 
margin ventrally, occiput with dense setig- 
erous punctures continuing from vertex, 
continuing on to gena. Gena in frontal view 
not swollen. Frons (above clypeus, below 
antennal sockets) slightly convex, with 
dense setigerous punctures nearly through- 
out, i.e., extending to clypeus and beyond 


VOLUME 102, NUMBER 4 


level of lower margin of compound eyes; 
with a central area immediately above clyp- 
eus smooth and glabrous, extending dorsal- 
ly % height of frons. Clypeus convex, pre- 
dominantly smooth and glabrous, dorsal 
margin marked by line of large setigerous 
punctures, hairs long and erect, margin 
broadly angled and raised medially, epis- 
tomal sulcus narrow and distinct medially, 
broad and poorly defined laterally. Malar 
space mostly punctate and pilose; malar 
sulcus present. Mandible with sparse setig- 
erous punctures on distignath, hairs elon- 
gate; gradually narrowed apically; biden- 
tate, teeth broadly rounded, ventral tooth 
larger, longer than dorsal; distignath broad- 
ly convex; subbasal groove fairly broad, 
margins rounded and somewhat indistinct, 
bottom of groove angled, appearing as a di- 
agonal sulcus extending from posterior 
mandibular articulation to lateral clypeal 
lobes; basignath distinctly widest near an- 
terior articulation, distignath reaching pos- 
terior articulation. 

Mesosoma: Pronotum predominantly 
smooth and glabrous dorsally; transition be- 
tween pronotal neck and collar broadly 
rounded in profile, marked by a weak trans- 
verse fold, laterally with sparse hairs and 
fine punctures. Mesoscutum predominantly 
smooth, very sparsely setose. Notauli ar- 
cuate, separated from transscutal articula- 
tion by short distance subequal to their 
width, without crenulae. Parapsidal furrows 
very indistinct. Scutellar pit wide, extend- 
ing laterally beyond notaulus, deep, without 
crenulae, crescentic, lateral margins of pit 
angled posteriorly; central convex portion 
of scutellum quadrate, separated from axilla 
by simple sulcus; posterior margin of scu- 
tellum with single transverse row of small 
foveae. Scutellum, axillae smooth, nearly 
glabrous. Dorsellum transverse, slightly 
bulging, anteriorly with a narrow crenulate 
sulcus, posteriorly mostly smooth. Meta- 
postnotum (anterior to transverse groove at 
base of propodeum) with pair of indistinct 
or broadly rounded posterior projections. 
Mesopleuron punctate and pilose nearly 


959 


throughout, save for smooth, glabrous, con- 
cave area adjacent to mesopleural suture. 
Mesepisternal groove indicated by a row of 
foveae extending ventrally from near base 
of fore wing to scrobal groove, then turning 
anteriorly and extending below fore coxa, 
widely separated from the discrimen ven- 
trally; scrobal groove indicated by trans- 
verse line of deep foveae; mesepisternum 
finely punctate and pilose, relatively flat 
ventrally, broadly rounded toward medial 
articulations, discrimen shallow, inconspic- 
uous, widened posteriorly to form small de- 
pression near coxae, with densely pilose, 
distinctly fingerlike lobe projecting above 
depression on each side. Metapleuron dis- 
tinctly separated from propodeum by row 
of deep, broad foveae; densely setose. Pro- 
podeum moderately globose in dorsal view, 
punctate along midline and around posterior 
end, dorsal surface otherwise smooth; dor- 
solaterally rugose, densely setose; anterior 
margin without teeth opposite metapostno- 
tal projections. Fore wing with radial cell 
closed, length 4.5/4.0 X width, base of m- 
cu only slightly displaced basad of bifur- 
cation of Cu,; in hind wing M between Cu, 
and Irm/Rs absent. 

Metasoma: First segment (petiole) dis- 
tinctly elongate, slender, strongly curved; 
remaining segments elongate, cylindrical, 
not laterally compressed; second segment 
longer than third, length of second 1.3/1.2 
length of third; second and third segments 
with tergite loosely wrapped around the 
sternite, not closely appressed and leaving 
very visible separation between tergum and 
sternum; length of metasomatic segments as 
a percentage of total length: 1: 23.6/23.5; 
2222319/22 42°32 18 3/1816; 4s 33: 
8.9/9.3; 6: 8.0/8.1; 7 to apex: 6.2/5.8. 

Male.—Other than characters of sexual 
dimorphism in metasoma and antenna typ- 
ical for family, differing from @ as follows: 
Body color generally brown above, yellow- 
ish brown below; base of legs including 
femora brownish yellow, brown apically. 
Fore wing length 4.2—4.9 mm (* = 4.54, 
SD = 0.268, n = 6). Sculpture on body 


960 


other than frons generally with punctures 
much less dense and largely smooth. Pro- 
podeum with sculpture along midline more 
extensive, extending from anterior to pos- 
terior margins. Antenna with dense short 
hairs, tyloid on A4—A8 small, with fine 
seta. Fore wing with closed radial cell, 
length 3.7—4.8 < width (* = 4.20, SD = 
0.379). Hind wing sometimes with a short 
stem of M arising from Irm/Rs. 

Material examined.—Holotype 2°: PAP- 
UA NEW GUINEA: Wau, 1,250 m, 
3.ix.1965, malaise trap, J.& M. Sedlacek 
(OSUC 117646, BPBM). Specimen in good 
condition; lacking A3—A15 from right an- 
tenna, A4—A15 from left. Paratypes: PAP- 
UA NEW GUINEA: (NE) Wau, Morobe 
Distr., 1,200 m, 5.x.1962, malaise trap, J. 
Sedlacek, 2 (OSUC 117653), 26.x.1961, 3 
(OSUC 117654), 23.x.1965, J. & M. Sed- 
lacek, 6 (OSUC 117647); (NE), Wau, Mo- 
robe Distr., 1,050 m, 11.ix.1961, malaise 
trap, J. Sedlacek, gd (OSUC 117651), 
30.1x.1961, ¢6 (OSUC 117652); ; NE Wau, 
Little Wau Ck., 1,200—1,300 m, 3.xii.1965, 
malaise trap, J. Sedlacek, 6 (OSUC 
117650); NE Karimui, 1,080 m, 14.vii.1963, 
M. Sedlacek (OSUC 117655). All speci- 
mens in BPBM. 

Etymology.—The specific epithet cra- 
cens, Latin for neat, slender, graceful, refers 
to the elongate petiole in the female of this 
species and its overall graceful habitus. 

Comments.—Monomachus cracens may 
be most easily distinguished from M. comp- 
tus by the remarkably long and curved pet- 
iole in the female and, in both sexes, the 
single row of foveae at the apex of the scu- 
tellum. The ventral lobes found at the pos- 
terior end of the mesepisternum are very 
distinct in male and female specimens; 
these lobes are visible only as weak raised 
margins of the depression in front of the 
mid coxae in M. comptus. Additionally, the 
lateral profile of the pronotum is mostly 
smooth and rounded, with only a weak line 
marking the separation of the neck and the 
collar dorsally. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Monomachus comptus Musetti and 
Johnson, new species 
(Figs. 2—3) 


Female.—Fore wing length 5.9 mm. Col- 
or: Head generally dark brown, clypeus 
and mandibles brown; lateral portions of 
pronotum, disc of mesoscutum on either 
side of notauli (including prescutum), pro- 
podeum, mesopleuron and metapleuron 
dark brown; scutellum, axillae, dorsal pron- 
otum, metanotum lighter in color, varying 
from brownish yellow to brown; metasoma 
brown; antenna brown; legs generally 
brownish yellow, hind leg beyond trochan- 
ter brown; wing membrane clear. 

Head: Transverse in frontal view (Fig. 
2), width across eyes 1.95 X wider than 
long, head width across the compound eyes 
(2.57 mm) slightly more than width across 
gena (2.46 mm). Eye height 3.13 malar 
length. Vertex and gena with same pattern 
of sculpture as found in M. cracens, but 
generally more smooth and shallow; outer 
margins of ocelli bordered by distinct fur- 
rows. Occipital carina short, widely sepa- 
rated from oral margin, occiput with dense 
setigerous punctures. Gena in frontal view 
not swollen. Frons flatter than M. cracens, 
with fine setigerous punctures extending be- 
low level of the compound eyes; central 
smooth, glabrous area above clypeus larger, 
mostly flat, extending through *% distance to 
antennal insertions; short raised carina pre- 
sent between antennae. Clypeus convex, 
smooth and nearly glabrous medially, shal- 
lowly punctate laterally, medially differen- 
tiated from frons by clear epistomal sulcus, 
without distinct line of large setigerous 
punctures, apical margin broadly angled 
and raised medially, epistomal sulcus poor- 
ly defined laterally. Malar area mostly 
smooth, with few sparse setigerous punc- 
tures near compound eyes, malar sulcus dis- 
tinct. Mandible with sparse setigerous 
punctures on distignath, hairs elongate; 
gradually narrowed apically; bidentate, 
teeth broadly rounded, ventral tooth wider, 
longer than dorsal; distignath broadly con- 


VOLUME 102, NUMBER 4 


961 


Figs. 2—4. 
matic segments 1-3, lateral view. 4, M. australicus, metasomatic segments 1—3, lateral view. Stippled lines on 
metasoma indicate lower margin of sterna visible through cuticle of terga. Scale line = 1 mm. 


vex; subbasal groove fairly broad, margins 
rounded and somewhat indistinct, bottom of 
groove angled, appearing as a diagonal sul- 
cus extending from posterior mandibular ar- 
ticulation to lateral clypeal lobes; basignath 
distinctly widest near anterior articulation, 
distignath reaching posterior articulation. 
Mesosoma: Pronotum predominantly 
smooth and glabrous dorsally; transition be- 
tween pronotal neck and collar abrupt, 
marked by ruga; ventrally with sparse hairs 
and fine punctures. Mesoscutum predomi- 
nantly smooth, very sparsely setose. Notau- 
li arcuate, separated from transscutal artic- 
ulation by short distance subequal to their 
width, finely crenulate. Parapsidal furrow 
present, extending over half length of me- 


Characters of Monomachus species. 2, M. comptus, head, frontal view. 3, M. comptus, metaso- 


soscutum from transscutal articulation. Scu- 
tellar pit slightly narrower, reaching later- 
ally as far as notaulus, with fine longitudi- 
nal crenulae, only slightly crescentic, lateral 
margins of pit broadly rounded; central 
convex portion of scutellum slightly widest 
anteriorly, separated from axilla by finely 
crenulate sulcus, posterior margin of scu- 
tellum with a subapical row of large punc- 
tures, apically with 2—3 rows of smaller fo- 
veae. Scutellum, axillae nearly smooth, gla- 
brous. Dorsellum less bulging than M. cra- 
cens, transverse, bordered anteriorly with 
sculptured sulcus half its length, posterior 
margin with narrow finely punctate band. 
Metapostnotum with pair of indistinct or 
broadly rounded posterior projections. Me- 


962 


sopleuron anteriorly punctate, setose ante- 
riorly and ventrally, with a wide smooth 
and glabrous area adjacent to mesopleural 
sulcus extending to intersection of scrobal 
and mesepisternal groove. Mesepisternal 
groove indicated by finely crenulate fold 
near wing base and by line of foveae ex- 
tending anteriorly from scrobal groove, 
these two sections widely separated, mes- 
episternal groove broadly separated from 
discrimen ventrally; scrobal groove indicat- 
ed by transverse line of deep foveae; mes- 
episternum finely punctate, sparsely setose, 
protuberant ventrally, closely abutting cox- 
ae so as to hide medial articulations, dis- 
crimen indicated by shallow longitudinal 
invagination, widened posteriorly to form 
small fusiform pit near mid coxae; margin 
of pit slightly produced and raised laterally. 
Metapleuron distinctly separated from pro- 
podeum by row of small foveae; densely 
setose. Propodeum moderately globose in 
dorsal view, coriaceous throughout, with 
scattered setigerous punctures, with vague 
indication of median longitudinal carina, 
posteriorly with longitudinal rugulae; dor- 
solaterally densely setose, longitudinally ru- 
gose; anterior margin without teeth opposite 
metapostnotal projections. Fore wing with 
radial cell closed, length 4.24 * width, base 
of m-cu only slightly displaced basad of bi- 
furcation of Cu,; in hind wing base of M 
present arising from Irm/Rs. 

Metasoma (Fig. 3): First segment slen- 
der, fairly straight; remaining segments 
elongate, cylindrical, not laterally com- 
pressed; length of second segment 1.3 X 
length of third; length of segments as a per- 
centage of total metasoma length: 1: 22.3; 
22 VOT 33: N83 As TAA Se 8; 62/947 
to apex: 6.7. 

Male.—Other than characters of sexual 
dimorphism in metasoma and antenna typ- 
ical for family, differing from female as fol- 
lows. Body color displaying same pattern 
as female except sometimes with less con- 
trast between dark brown and brownish yel- 
low areas. Fore wing length 4.8—6.2 mm (x 
= 5.46, SD = 0.622, n = 4). Vertex with 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


more extensive smooth areas. Propodeum 
with spiracle bulging, dorsolaterally some- 
times with well defined longitudinal carina. 
Antenna with dense short hairs, tyloid on 
A4—A8 small, with fine erect setae. Fore 
wing with radial cell closed, length 3.1—4.0 
x width (« = 3.75, SD = 0.295). Hind 
wing sometimes with M absent between 
Cu, and Irm/RS. 

Material examined.—Holotype 2: PAP- 
UA NEW GUINEA: SE Mount Giluwe, 
2,500—2,750 m, 30.v.1963, J. Sedlacek 
(OSUC 117648, BPBM). Specimen in fair- 
ly good condition; lacking A5—A15 from 
right antenna, and A1l3—A15 from left. Ab- 
domen broken, segments four and beyond 
in gelatin capsule attached to pin. Paraty- 
pes: PAPUA NEW GUINEA: Daulo Pass, 
2,450 m, 22.xii.1978-8.1.1979, J. Sedlacek, 
6 (OSUC 117079, AEIC); Mount Otto, 
2,000 m, 22.xii.1978-9.1.1979, J. Sedlacek, 
6 (OSUC 117077, AEIC); Tari Gap, nr. 
Mount Hagen, 2,600 m, 29.i-4.11.1979, J. 
Sedlacek, 6 (OSUC. 117080; “AEBI@); 
Mount Giluwe, 2,800 m, 3.1-8.11.1979, J. 
Sedlacek, ¢ (OSUC 117078, AEIC). 

Etymology.—The specific epithet comp- 
tus, Latin for ornamented, refers to the 
more elaborate sculpture on the scutellum. 

Comments.—See diagnosis under de- 
scription of Monomachus cracens for the 
most useful characters to distinguish the 
two Papuan species. 


DISCUSSION 


Schulz (1911) distinguished the New 
World species of Monomachus from those 
in Australia by means of the curvature of 
the petiole: strongly bowed in specimens 
from America, straight in those from Aus- 
tralia. Monomachus cracens (Fig. 1) clearly 
does not conform to this rule. Both species 
from New Guinea are very similar to M. 
australicus Girault in terms of the structure 
of the mandibles, the sculpture on the body, 
the shape of the head, and the shape of the 
clypeal margin. The color patterns observed 
differ from that ‘‘yellow form’ described 
by Naumann for specimens of M. australi- 


VOLUME 102, NUMBER 4 


cus from northern Queensland most notice- 
ably in that the lateral lobes of the meso- 
scutum are the same color, dark brown, as 
the medial lobe. The ventral mesepisternal 
lobes are also shared with all Australian 
Monomachus. The Australian species are 
immediately distinguishable on the basis of 
the wing venation: the base of m-cu is 
strongly displaced basad of the bifurcation 
of Cu, in the fore wing. The vein is inserted 
nearly in the middle of cell 2Cu (first sub- 
discal cell). All of these Old World species 
have the second and third metasomatic seg- 
ments elongate, with the second longer than 
the third. Neotropical Monomachus typical- 
ly have the second segment much shorter 
and apically widened, and the metasoma 
beyond the petiole is laterally compressed. 


ACKNOWLEDGMENTS 


Thanks to D. Wahl (AEIC), G. Nishida 
(BPBM), L. Masner (Ottawa), S. Lewis 
(London), P. Perkins (Cambridge), and J. C. 
Cardale (Canberra) for the loans of speci- 


963 


mens. This material is based upon work 
supported by the National Science Foun- 
dation under Grant No. DEB-9521648. 


LITERATURE CITED 


Huber, J. T. and M. J. Sharkey. Structure, pp. 13-59. 
In H. Goulet and J. T. Huber, eds., Hymenoptera 
of the World: An Identification Guide to Families. 
Agriculture Canada. Research Branch. Publication 
1894/E. Ottawa, Canada, 668 pp. 

Johnson, N. E and L. Musetti. In press. Data ware- 
housing architecture and tools for Hymenoptera 
biodiversity informatics. Jn Austin, A. D., ed., 
Proceedings of the 4 International Hymenoptera 
Conference. Canberra, Australia. 

Masner, L. 1993. Superfamily Proctotrupoidea, pp. 
537-557. In Goulet, H. and J. T. Huber, eds., Hy- 
menoptera of the World: An Identification Guide 
to Families. Agriculture Canada. Research 
Branch. Publication 1894/E. Ottawa, Canada, 668 
Pp- 

Naumann, I. D. 1985. The Australian species of Mon- 
omachidae (Hymenoptera: Proctotrupoidea), with 
a revised diagnosis of the family. Journal of the 
Australian Entomological Society 24: 261-274. 

Schulz, W. A. 1911. Systematische Uebersicht der 
Monomachiden. Memoirs of the International 
Congress of Entomology 2: 405—422. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 964-968 


DESCRIPTION OF THE PUPA OF ARMIGERES (LEICESTERIA) OMISSUS 
(EDWARDS) AND A KEY TO THE LARVAE AND PUPAE OF THE 
ARMIGERES OCCURRING IN NEPAL (DIPTERA: CULICIDAE) 


RICHARD FE. DARSIE, JR. 


Florida Medical Entomology Laboratory, University of Florida, 200 9th Street SE, Vero 
Beach, FL 32962, U.S.A. (e-mail: rfd@ gnv.ifas.ufl.edu) 


Abstract.—The pupa of Armigeres omissus (Edwards) is described for the first time. 
Keys to the larvae and pupae of Armigeres species occurring in Nepal are added. 


Key Words: 


The pupal stage of species of the sub- 
genus Leicesteria, genus Armigeres occur- 
ring in Nepal were described by Darsie 
(1998). Subsequently, a female of Armiger- 
es (Leicesteria) omissus (Edwards) was dis- 
covered in the collection of Nepal mosqui- 
toes at the Florida Medical Entomology 
Laboratory, Vero Beach, FL, a new country 
record. Its pupa was briefly described by 
Delfinado (1966) and Baisas (1974). A 
more detailed description follows, based on 
specimens from Thailand since none are 
available from Nepal. 

With this account, the pupae of all spe- 
cies of Armigeres from Nepal have now 
been described (Ramalingam 1987; Toma et 
al. 1994; Darsie 1998, 2000). Inasmuch as 
a detailed study of the larvae was a neces- 
sary part of the pupal verification, a larval 
key is also included. This is a revision of a 
key by Darsie and Pradhan (1990). Since 
then, four species have been added to the 
fauna, i.e., Ar. (Arm.) theobaldi Barraud 
(Pradhan and Darsie 1990), Ar. (Lei.) in- 
choatus Barraud and Ar. (Lel.) digitatus 
(Edwards) (Darsie et al. 1992) and Ar. 
(Lei.) omissus, herein. 

The pupa of Ar. omissus possesses the 
generic and subgeneric characters given be 
Darsie (2000). It is readily distinguished 


Armigeres omissus, pupa, keys, Nepal 


from pupae of the Nepal Armigeres species 
by a combination of: the absence of seta 1- 
P, the paddle fringe extending to near the 
base, seta 6-II-V with thin branches and 
seta 1-II with 17 or fewer branches. 


METHODS AND MATERIALS 


For procedures used in this study refer to 
Darsie (1998). No pupae of Ar. omissus 
were found in my collection from Nepal, 
but specimens were borrowed from the 
Walter Reed Biosystematic Unit, National 
Museum of Natural History (NMNH), 
Smithsonian Institution, with accompany- 
ing larval exuviae for species verification. 
In the description below br means branches 
and Le and Pe mean exuviae of the fourth 
instar larva and pupa, respectively. 


DESCRIPTION 


Armigeres (Leicesteria) omissus (Edwards) 
(Fig. 1) 

Position and size of setae as figured, 
range and modal number of branches in Ta- 
ble 1. Cephalothorax: Setae 1,3-CT long to 
very long, thin, usually single (1,2); 6-CT 
0.53—-0.82, * 0.66 length of 7-CT; trumpet 
brown, reticulate, length 0.5—0.6 mm, index 
1.54-2.50, x 2.05. Abdomen: Seta 1-Il 
moderately long, with 10—17 br; 2-V-VII 


VOLUME 102, NUMBER 4 965 


PRorlde Toye 


Fig. 1. Pupa of Armigeres (Lei.) omissus. A, Cephalothorax (left side). B, Metanotum and abdomen (dorsal 
left, ventral right). Abbreviations: CT = cephalothorax; GL = genital lobe; Mr = paddle midrib; Pa = paddle; 
T = respiratory trumpet. 


966 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
Table 1. Pupa chaetotaxy of Armigeres omissus. 
Abdominal segments 
Cephalo 

Seta thorax I II II IV Vv VI Vil Vill 
0) - = 1 1 1 1 l 

l i 2DY 5-10 (?) 10-17 (14) 2-8 (3) 25 (3) 2-6 (2) 1—3 (2) De 2) (C2) - 

2 1 (2) 1 | l 1 1 1 _ 

3 ey 2D} l 1 N=7/ (6) =s)(G)) 1-3 (1) 27a) = 

4 Don) 24 (3) 2-6 (4) 1-3 (1) IES Gls) 122) 1-3 (1) 1, 2 (1) We 22, ((Ib)) 
5 1-3 (2) l 2-5 (2) 1-5 (3) 1-3 (1) ee Gy) 122) 2 (GD) - 

6 ] 1—4 (?) 2-7 (2) 35 (3) 14 (2) ils ZA) ls 2 (Ub) 14 (1) _ 

7 12119) 1 2@)* 2-5: (2)" 2, 32) 133) 24@€4) 1-3 (2) 1-3 (1) - 

8 1-3 (2) = = BIC) BrayQy 2x 3@)y 220 @) 2-8 (?) 2-5 (5) 
9 1 ] 1 1 1 1 Is) (2) = 
10 1 (2) - 12) 2n@) 1-3 (2) 1-3 (1) 1 1 _ 

11 1 - = 1—3 (1) 1-3 (1) 1-3 (1) 1-3 (1) 1-3 (2) - 
12 1 - - - - ~ = - ~ 
14 _ — - 1 1 l 1 tS 7 (Ob) 1 


' Range followed in parenthesis by mode. 


short, single, 0.22—0.40, « 3.0 length of 1- 
V-VII; 3-ILIII stout, 0.62-0.91, x 0.75 
length of following tergum; 5-IV,V stout, 
more than 0.5 length of following tergum; 
6-IJ]-V short, with thin branches, usually 
double (1-5); 6-VI long, stout, sparsely 
aciculate, single, seldom double; 9-VII 
long, stout, aciculate, with 2—5 br, seldom 
single; 9-VIII long, stout, aciculate, usually 
with 5 br (2-5). Seta 12-IV was found in 
only one pupa. Paddle: Length 1.10—1.27 
mm, index 1.05—1.29, 1-P absent, except 
one pupa with a seta-like spicule without 
alveolus, fringe long, 0.19 mm, extending 
to near base. 

The description is based on the follow- 
ing specimens, all from Thailand, depos- 
ited in the NMNH: Chiang Mai Province, 
Huey Muang Ban Ay, V-15-64, 12 LePe, 
ex bamboo pot; Nan Province, Ban Pha 
Hang, elev. 400 m, VIII-19-66, 22 LePe, 
ex bamboo stump; Lampang Province, Doi 
Pha Huat, elev. 420 m, V-21-68, 12 LePe, 
ex bamboo stump. The Nepal specimen 
was collected in Jhapa District, Kanchan- 
bari, VIII-2-91, 1°, resting outdoors on 
vegetation in primary forest (coll. no. 111- 
x28): 


KEY TO PUPAE OF ARMIGERES SPECIES OF 
NEPAL 


Ihe Seta 1-P small or absent; seta 5-IV-V at 
least 0.5 length of following tergum; seta 
6-CT shorter than 7-CT (subg. Leicester- 


TGV. bbe odode ors eee 2 
- Seta 1-P long, rather stout, if not, seta 5- 
IV,V less than 0.5 length of following ter- 
gum (theobaldi); seta 6-CT as long as or 
longer than 7-CT (subg. Armigeres) 8 
2(1) Paddle with fringe of long spicules on 
outer margin extending to near base ... 3 
~ Paddle with fringe of long spicules on 
outer margin in apical 0.75 or less .... 6 
3(2) Seta 6-II-V short, usually with 2 or more 
thin branches’). «2 32.55 See eee 4 
= Seta 6-II-V long, stout, single ....... 3 
4(3) Seta 1-II with 28 or more branches; seta 
1-I with thick unbranched base, 0.27 of 
totallenctheepses eee digitatus (Edwards) 
— Seta 1-II with 17 or fewer branches; seta 
1-I with smaller unbranched base, 0.18 of 
totaljlenothiesress eee omissus (Edwards) 
5(3). Seta 3-IV usually 5- or 6-branched; seta 
I= sparsely aciculate == 0-3-1) 
SPS eM ees aca ahs, ‘ool OZ annulitarsis (Leicester) 
— Seta 3-IV usually with 4 or fewer branch- 
es; seta 1-II densely aciculate ...... 
aaah) OTE en fy Witte Veg Z wellh magnus (Theobald) 
6(2). Seta 1-II with 24 or more branches; pad- 


dle with large external lobe; seta 1-II-VII 
subequal toxsetal 2a oe 
dolichocephalus (Leicester) 


VOLUME 102, NUMBER 4 


- Seta 1-II with 21 or fewer branches; pad- 
dle without large external lobe; seta 1 at 
least 2.0 length of seta 2 on V-VII.... 7 
Seta 3-CT with thin branches; seta 6-VI 
single inchoatus Barraud 
- Seta 3-CT with stout branches; seta 6-VI 
usually with 2 or more branches ... . 
sic Joust SD DESORBED TOR TLCe SP eRe dentatus Barraud 
Setae 3-II, III and 5-IV, V shorter than 
0.25 length of following tergum . . 
theobaldi Barraud 
- Setae 3-II, III and 5-IV, V longer than 0.5 
length of following tergum 
Seta 1-CT with thin branches, usually 
double; seta 9-VII with 9 or more branch- 
CS eee ae eee ICE. Pe aureolineatus (Leicester) 
- Seta 1-CT stout, single or double; seta 9- 
VII with 8 or fewer branches ....... 10 
Seta 1-II small, usually with 5 or fewer 
thin branches, 0.36 or less length of 3-II 
od cages en ot DED U EERE kuchingensis Edwards 
- Seta 1-II much thicker, with 6 or more 
branches, 0.5 or more length of 3-II .. 11 
1110). Seta 1-II pedunculate, brush-like, 
branched in apical 0.7, with with 17 or 
more branches ........ durhami Edwards 
- Seta 1-II pedunculate or not, with 6-14 
branches 
12(10). Trumpet short and broad, index 1.2—1.7: 
seta 3-VII closer to seta 4 than to seta 1 
Seer a ts oes ciin subalbatus (Coquillett) 
- Trumpet long, index 2.3 or greater; seta 
3-VII closer to seta | than to 4 


7(6). 


8(1). 


9(8). 


10(9). 


KEY TO FOURTH INSTAR LARVAE ARMIGERES 
SPECIES OF NEPAL 


(Partially adapted from Macdonald 1960) 


1. Abdominal segment X with dorsal saddle 
and very small ventral sclerotized plate 
Sie Sask eae eee ne magnus (Theobald) 
= Abdominal segment X with dorsal scler- 


alized saddle Onllys .\3. ots She annie acs 2 
2(1) Comb scales fringed with subequal spi- 
TUES. ta i eatin Uae Sy ee ae at ae 3 
— At least some comb scales with apical 
SJOVCIYES | onrn pe ery GEE SARC Phan ee Aaah eee eee 8 
3(2) Comb with more than 25 scales 
55: 5 6 a OND ORO ROR eRe: inchoatus Barraud 
= Comb with fewer than 25 scales...... 4 
4(3) Comb with 18=25 scales =~... 4 62. 
A Mate ke, Be ES Re annulitarsis (Leicester) 
= Comb with fewer than 18 scales...... 5 
5(4). Most comb scales rather pointed apically, 
point fringed with subequal spinules ... 6 


= Comb scales rounded apically, fringed 
with subequal spinules.............. 7 


967 
6(5). Seta 1-X on saddle or close to it ..... 
Bede Tea ae Ea durhami Edwards 
= Seta 1-X distinctly removed from saddle 
BS PERS ROMEO fo “opis subalbatus (Coquillett) 
7(5) Seta 1-III-VI reaching posterior margin of 
following segment; seta 6-I with 3 or 4 
[RINENES, 9.666 cielo + kuchingensis Edwards 
- Seta 1-III-VI only reaching basal 0.25 or 
less of following segment; seta 6-I with 
D9) MONS. 255-55. kesseli Ramalingam 
8(2) Comb with 11 or fewer scales ....... 9 
= Comb with 12 or more scales....... 10 
9(8) Seta 1-S very small, with 2 or 3 branches; 
1-X small, not inserted on saddle ..... 
SOY ree eee aureolineatus (Leicester) 
= Seta 1-S long, rather stout, single; seta 1- 
X strong, inserted on saddle ....... 
Ny ae.’ bee syn cee oy. ER theobaldi Barraud 
10(9). Abdominal segments I-VII with promi- 
nent tubercles bearing setae ....... 
Bs gas ea isie sod cae 6 dolichocephalus (Leicester) 
~ Abdominal segments I-VII without tu- 
bercles ws: 4: Aaa vats a eee oes i 
11(10). At least abdominal sterna I-IV with 


large patch of fine spicules ....... 
digitatus (Edwards) 
= Abdominal sterna I-IV without spicules 


12(11). Seta 5-VIII with 3 or more fine branches: 
comb scales with 2 or more apical spines 

ae es sence ae omissus (Edwards) 
= Seta 5-VIII stout, single or double; most 
comb scales with single apical spine . . . 

SERS s RR RY chanetined open dicws dentatus Barraud 


ACKNOWLEDGMENTS 


The author is indebted to the National 
Geographic Society for their support of the 
work, to T. Gaffigan, Walter Reed Biosys- 
tematics Unit, for providing the specimens 
used in this study, to B. Bower-Dennis for 
the illustration, and to J. R. Rey and J. K. 
Nayar for reviewing the manuscript. This is 
Florida Agricultural Experiment Station 
Journal Series No. R-06543. 


LITERATURE CITED 


Baisas, F E. 1974. The mosquito fauna of Subic Bay 
Naval Reservation, Republic of the Philippines. 
U.S. Navy, Headquarters First Medical Service 
Wing Technical Report 72-2, 1-170. 

Darsie, R. F, Jr. 1998. Descriptions of the pupae of six 


968 


species of Armigeres Theobald, subgenus Leices- 

teria Theobald (Diptera: Culicidae) from Nepal. 

Proceedings of the Entomological Society of 

Washington 100: 234—246. 

. 2000. Description of the pupae of five species 
in subgenus Armigeres Theobald, genus Armiger- 
es Theobald, with a key to species of the known 
pupae of the subgenus (Diptera, Culicidae). Pro- 
ceedings of the Entomological Society of Wash- 
ington 102: 108-119. 

Darsie, R. F, Jr. and S. P. Pradhan. 1990. The mos- 
quitoes of Nepal their identification, distribution 
and biology. Mosquito Systematics 22: 69-130. 

Darsie, R. F, Jr., S. P. Pradhan, and R. G. Vaidya. 1992. 
New species records from 1991 collections. Mos- 
quito Systematics 24: 23-28. 

Delfinado, M. D. 1966. The culicine mosquitoes of the 
Philippines, tribe Culicini (Diptera, Culicidae). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Memoirs of the American Entomological Institute 
(Ann Arbor) 7: 1—252. 

Macdonald, W. W. 1960. Malaysian parasites 
XXXVIII. On the systematics and ecology of Ar- 
migeres subgenus Leicesteria (Diptera, Culicidae). 
Studies of the Institute for Medical Research 29: 
110-153. 

Pradhan, S. P. and R. FE Darsie, Jr. 1990. New additions 
to the mosquito fauna of Nepal. Journal of the 
Institute of Medicine (Nepal) 12: 225-228. 

Ramalingam, S. 1987. On the restriction of Armigeres 
durhami Edwards and the description of Armiger- 
es kesseli n.sp. (Diptera: Culicidae). Tropical Bio- 
medicine 4: 55—65. 

Toma, T., I. Miyagi, and N. Benjaphong. 1994. Rede- 
scription of Armigeres (Armigeres) theobaldi 
(Diptera: Culicidae). Mosquito Systematics 26: 
11-18. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 969-973 


NEW RECORDS OF FLEAS (SIPHONAPTERA) FROM 
EASTERN WEST VIRGINIA 


RALPH P. ECKERLIN AND HARRY EF PAINTER 


Natural Sciences Division, Northern Virginia Community College, Annandale, VA 


22003 U.S.A. (e-mail: reckerlin@nv.cc.va.us) 


Abstract.—Nine species of fleas are reported for the first time from the boreal envi- 
ronment in the mountains of eastern West Virginia. The new state records are Cerato- 
phyllus gallinae (Schrank), Conorhinopsylla stanfordi Stewart, Epitedia faceta (Roths- 
child), Hystrichopsylla tahavuana Jordan, Megabothris asio asio (Baker), M. quirini 
(Rothschild), Nearctopsylla genalis genalis (Baker), Rhadinopsylla media Smit, and Tam- 


iophila grandis (Rothschild). 
Key Words: 


We report nine flea species found in the 
higher elevations of the Appalachian Moun- 
tains of West Virginia along the eastern bor- 
der with Virginia where boreal conditions 
are found. There has been considerable 
study of the biota of the Appalachian balds 
(Mark 1958) and spruce and northern hard- 
wood forests (Handley 1971, Stephenson 
and Adams 1984, and references therein) 
demonstrating plants and animals charac- 
teristic of the boreal environments of more 
northern regions. Results of these studies 
suggested to us that fleas common to more 
northern environments might be present in 
West Virginia because of the presence of 
appropriate mammal hosts and the exis- 
tence of boreal conditions suitable for their 
non-parasitic larval stages. Benton (1971, 
1980) added seven new species to the West 
Virginia list bringing the state total to 24 
and further noted that an additional 20 fleas 
not then known from the state should be 
present. We add nine new flea species to the 
West Virginia list with all but one expected 
by Benton. 


Appalachian Mountains, fleas, distribution, West Virginia 


MATERIALS AND METHODS 


Collections were made in the eastern 
mountains of the Ridge and Valley Province 
of West Virginia as often as could be ar- 
ranged during all months of the year from 
1984 to 1996. Bird and mammal nests were 
placed in a Berlese funnel overnight in the 
laboratory to extract fleas. Small mammals 
were trapped using Sherman live traps and 
snap traps. Live animals were placed in pa- 
per bags containing paradichloro-benzene 
crystals for a few minutes and then released 
into plastic bags for identification, sexing 
and a check for fleas still in the pelage. The 
animals were then released in the vicinity 
of their capture. Collection data were writ- 
ten on the paper bag and the contents were 
examined for fleas in the laboratory. Mam- 
mals found dead were brushed over a white 
enameled pan and fleas were placed in eth- 
anol. Five nest boxes were placed about 3 
m high in a stand of red spruce (Picea rub- 
ens Sargent) on Alleghany Mountain in Po- 
cahontas County at an elevation of 1,300 m 
to monitor the fleas of flying squirrels. 


970 


White polyester fiber was provided as nest- 
ing material and sampled as nearly monthly 
as weather would permit for a year. Sam- 
pling involved removal of only half of the 
fiber from a nest box, placing it in a plastic 
bag and replacing it with fresh fiber, thereby 
permitting the maintenance of succeeding 
generations of parasites. Fleas were re- 
moved from the retained nesting material in 
the lab and stored in ethanol. All fleas were 
decolorized in 10% KOH, dehydrated in 
ethanol, cleared in xylene and mounted in 
Canada balsam. Fleas were identified using 
keys developed by Benton (1983) and Hol- 
land (1985). Mammal names follow Wilson 
and Reeder (1993). 


RESULTS 


The following nine flea species new to 
the state of West Virginia were collected. 


Ceratophyllidae 
Ceratophyllus gallinae (Schrank) 


3 2 from the nest of Sialia sialis (L.), 26 
Sept 1993, Pocahontas County, Top of Al- 
legheny. 

This European hen flea is a wide-ranging 
species from Alaska and Canada (Holland 
1985) as well as northeastern United States 
(Benton 1980). Domestic chickens and na- 
tive birds are the usual hosts. This is the 
southernmost reported occurrence of this 
species in the eastern United States. 


Megabothris quirini (Rothschild) 


1 2 ex Clethrionomys gapperi (Vigors), 
15 Nov 1984, 1 2 ex C. gapperi, 20 April 
1985, 1 2 ex Microtus pennsylvanicus 26 
Sept 1993, all from Pocahontas County, Al- 
leghany Mountain. 1 6, 2 2 ex C. gapperi 
30 June 1985, 1 ex Peromyscus leucopus 
(Rafinesque), 30 June 1985, Randolph 
County, Spruce Knob Lake. 4 2 ex C. gap- 
peri, 29 Aug 1985, 1 3 ex M. chrotorrhinus 
(Miller), 9 Dec 1985, 3 2 ex C. gapperi 18 
May 1986, all from Pendleton County, 
Spruce Knob. 

Hopla (1965) stated that M. quirini “‘is a 
flea that has migrated into the boreal re- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


gions from the south following the retreat 
of the Pleistocene glaciers.” It is found on 
mice in seven genera (Holland 1985). Our 
collections were primarily from microtines. 
Five of 72 C. gapperi, 1 of 9 M. chrotor- 
rhinus and 1 of 27 P. leucopus were in- 
fested. In the eastern United States this flea 
had been reported only from New York and 
New England (Benton 1980). The nearest 
record to those reported here is from south- 
ern New York, over 500 km to the north- 
east. 


Megabothris asio asio (Baker) 


1 3, 26 2 ex 9 Microtus pennsylvanicus 
and 1 6 ex Peromyscus maniculatus (Wag- 
ner), 18 Nov 1984; Pocahontas County, Al- 
leghany Mountain; 2 2 ex 2 M. pennsyl- 
vanicus, 1 3 ex P. maniculatus and 1 3 ex 
C. gapperi, 20 Oct 1985 from Tucker 
County, Dolly Sods; 1 6 ex M. chrotor- 
rhinus, 9 Dec 1985 Pendleton County, 
Spruce Knob; 1 6, 4 2 ex M. pennsylvan- 
icus 30 Jun 1985, Randolph County, Spruce 
Knob Lake. 

Numerous collections were made at these 
four sites during the Fall, Winter and Spring 
months. This is another flea found primarily 
on microtine hosts with a boreal distribu- 
tion in eastern Canada and northeastern 
United States. The previous nearest record 
of this flea is in Pennsylvania. Thus, our 
Appalachian records from West Virginia 
extend the range about 300 km to the south. 


Ctenophthalmidae 
Conorhinopsylla stanfordi Stewart 


1 36,1 2 ex Glaucomys volans (L.) nest, 
10 Nov. 1985, Pocahontas County, Alle- 
ghany Mountain. 

This flea is blind and is primarily a nest 
flea on flying squirrels. Other hosts include 
squirrels in the genera Sciurus and Tamias- 
ciurus (Holland 1985). It is known from 
Ontario to Maryland so this is the south- 
ernmost occurrence in the East. This flea is 
not plentiful in collections, probably due to 
a dearth of nest examinations of the flying 
squirrel in winter. It was found in larger 


VOLUME 102, NUMBER 4 


numbers in more extensive nestbox arrays 
in neighboring Virginia. 


Tamiophila grandis (Rothschild) 


2 2 ex Tamias striatus (L.), 25 Sep 1993, 
12 Oct 1996, Pocahontas County, Top of 
Allegheny. 

As the name implies, this is a large flea 
and a parasite of the chipmunk, of which 2 
of 15 were parasitized. This flea is rare in 
collections because it is a nest flea and is 
rarely taken on the host. It occurs from On- 
tario to southwestern Virginia where it was 
recently reported by Eckerlin and Painter 
01995): 


Epitedia faceta (Rothschild) 


2 2 from nest of G. volans, 19 Jan 1986, 
Pocahontas County, Alleghany Mountain. 

This nest flea is most abundant in the late 
fall and early winter (Benton 1980). Both 
northern and southern flying squirrels are 
hosts but the flea is restricted to eastern 
Canada and eastern United States, corre- 
sponding closely with the range of G. vo- 
lans (Holland 1985). The southernmost re- 
ported occurrence of E. faceta is eastern 
Tennessee in the Appalachians at high ele- 
vation (Durden and Kollars 1997). 


Nearctopsylla genalis genalis (Baker) 


3 2 ex B. brevicauda, 9 Dec 1985, Pen- 
dleton County, Spruce Knob; 1 @ ex B. 
brevicauda, 3 Nov 1995, Pocahontas Coun- 
ty, Alleghany Mountain, 4 2 ex Sorex fu- 
meus Miller, 18 Nov 1984, Pocahontas 
County, Top of Allegheny. 

Benton (1980) refers to this flea as a fall 
and winter flea and indeed our records are 
all in the fall months. With a range from 
Ontario and Illinois eastward to the Atlantic 
seaboard states, N. genalis was previously 
collected no further south than Pennsylva- 
nia. The Pocahontas County site extends the 
range of this species 300 km southward at 
high elevations in the Appalachians. Two of 
33 B. brevicauda and 1 of 2 S. fumeus were 
infested. 


O71 


Rhadinopsylla media Smit 


1 2 from nest of Neotoma floridana 
(Ord), 17 Sep 1950, Pendleton County, 
Franklin, collected by Price and Tipton. 

The single damaged specimen is depos- 
ited in the Monte L. Bean Life Sciences 
Museum at Brigham Young University and 
was kindly loaned us by Michael Hastriter. 


Hystrichopsyllidae 
Hystrichopsylla tahavuana Jordan 


1 @ ex Blarina brevicauda (Say), 18 
May 1986, Randolph County, Spruce Knob 
Lake, and 1 2 from nest of Microtus penn- 
sylvanicus (Ord), 25 Oct 1986, Pocahontas 
County, Top of Allegheny. 

This large flea was described from the 
two host species we list. Scattered records 
exist from Ontario to Tennessee. Only 1 of 
33 B. brevicauda was infested so we regard 
this as an uncommon flea. 


DISCUSSION 


Eight of the nine new records are species 
Benton (1971) predicted would be found in 
West Virginia. The only unexpected species 
was Rhadinopsylla media whose nearest re- 
ported occurrence is in Michigan and Min- 
nesota (Benton and Timm 1976). Addition- 
al comment is warranted by the consider- 
ably greater physical separation between 
this record and other specimens of this sel- 
dom collected flea. Pfitzer (1950), in his un- 
published master’s degree thesis reported 
collecting a number of the closely related 
Rhadinopsylla orama Smit 1957 from Wise 
County, Virginia just north of the Tennes- 
see/Virginia line. The number and location 
of genal spines (of the R. media specimen, 
Fig. 1) confirm placement of the West Vir- 
ginia specimen in Rhadinopsylla and the 
long suture between the metanotum and 
metepimeron (Fig. 2) separates R. media 
from the only other eastern species, R. or- 
ama, which is found in the adjacent states 
of Virginia, Pennsylvania, and Maryland 
(Benton 1980), but was not observed in this 
study. We agree with Benton and Timm 


972 


Figs. 1-2 
eron. 


Rhadinopsylla media. 1, Head. 2 


(1976) that more specimens are required to 
determine distribution of these poorly 
known fleas. 

Epitedia faceta, Hystrichopsylla tahavu- 
ana, Megabothris a. asio, M. quirini, 
Nearctopsylla g. genalis and Tamiophila 
grandis have extensive distributions north 
of West Virginia, but are restricted to high 
elevations southward in the Appalachian 
mountains. We regard these species and 
their environments in West Virginia as bo- 
real. Significant extensions of ranges were 
demonstrated for N. g. genalis and M. a. 
asio, previously not collected south of 
Pennsylvania and M. quirini, not collected 
south of New York. Conversely, Conorhin- 
opsylla stanfordi and Ceratophyllus galli- 
nae are widespread and not ecologically 
bound to the Appalachians. Conorhinopsyl- 
la stanfordi adults in the nests of flying 
squirrels have a pronounced cold weather 
peak of abundance and are absent for most 
of the year (Benton and Day 1980). Many 
species of birds serve as hosts of C. galli- 
nae including many cavity nesters such as 
bluebirds (Holland 1985) which are found 
throughout the study area. The total number 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


“))))))) yo 


, Metathorax showing suture between metanotum and metepim- 


of species reported from West Virginia now 
stands at 33. We believe that more of Ben- 
ton’s (1971) hypothesized species will be 
discovered in West Virginia with additional 
collecting. 


ACKNOWLEDGMENTS 


We acknowledge with gratitude the tu- 
telage and help of the late Robert Traub in 
our studies. He made records available to 
us, permitted us use of his reference collec- 
tion and literature, taught us flea anatomy, 
and constantly urged us to get out and do 
more collecting. We thank the West Virgin- 
ia Department of Natural Resources for 
granting collection permits. We thank the 
many students who helped us collect or 
who donated specimens. Finally, we thank 
the reviewers for comments and sugges- 
tions that have greatly improved this paper. 


LITERATURE CITED 


Benton, A. H. 1971. An annotated list of the fleas (Si- 
phonaptera) of West Virginia. Proceedings of the 
West Virginia Academy of Science 40: 35-39. 

. 1980. An atlas of the fleas of the eastern Unit- 

ed States. Marginal Media, Fredonia, New York, 

177 pp. 


VOLUME 102, NUMBER 4 


. 1983. An illustrated key to the fleas of the 
eastern United States. Marginal Media, Fredonia, 
New York, 34 pp. 

Benton, A. H. and J. F Day. 1980. Seasonal changes 
in the flea fauna of nests of the southern flying 
squirrel, pp. 401—403. Jn Traub, R. and H. Starke, 
eds., Fleas. Proceedings of the International Con- 
ference on Fleas, Ashton Wold, Petersborough, 
UK 21-25 June, 1977. A. A. Balkema, Rotterdam 

Benton, A. H. and R. M. Timm. 1976. Description of 
the female of Rhadinopsylla media Smit (Siphon- 
aptera: Hystrichopsyllidae). Journal of Medical 
Entomology 13: 473—475. 

Durden, L. A. and T. M. Kollars, Jr. 1997. The fleas 
(Siphonaptera) of Tennessee. Journal of Vector 
Ecology 22: 13-22. 

Eckerlin, R. P. and H. E Painter. 1995. First record of 
Tamiophila grandis (Insecta: Siphonaptera) from 
Virginia. Banisteria No. 6: 24—25. 

Handley, C.O., Jr. 1971. Appalachian mammalian ge- 
ography—Recent epoch, pp. 263-303. Jn Holt, P. 
C., R. A. Paterson, and J. P. Hubbard, eds., The 
Distributional History of the Biota of the Southern 
Appalachians Part III. Vertebrates. Research Di- 


973 


vision Monograph 4, Virginia Polytechnic Insti- 
tute and State University, Blacksburg, Virginia. 

Holland, G. W. 1985. The fleas of Canada, Alaska and 
Greenland (Siphonaptera). Memoirs of the Ento- 
mological Society of Canada No. 130, Entomo- 
logical Society, Ottawa, Canada, 631 pp. 

Hopla, C. E. 1965. Alaskan hematophagous insects, 
their feeding habits and potential as vectors of 
pathogenic organisms. I. The Siphonaptera of 
Alaska. Arctic Aeromedical Laboratory, Fort 
Wainwright, Alaska, 267 pp. 

Mark, A. E 1958. The ecology of the southern Appa- 
lachian grass balds. Ecological Monographs 28: 
293-336. 

Pfitzer, D. W. 1950. A manual of the fleas of Tennes- 
see, M.S. Thesis, University of Tennessee, Knox- 
ville, 103 pp. 

Stephenson, S. L. and H. S. Adams. 1984. The spruce- 
fir forest on the summit of Mount Rogers in south- 
western Virginia. Bulletin of the Torrey Botanical 
Club 111: 69-75. 

Wilson, D. E. and D. M. Reeder, eds. 1993. Mammal 
species of the world, A taxonomic and geographic 
reference, 2d Edition., Smithsonian Institution 
Press, Washington, D.C., 1206 pp. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 974-990 


SAWFLIES (HYMENOPTERA: TENTHREDINIDAE) DESCRIBED BY 
BENJAMIN D. WALSH, WITH NOTES ON THEIR HOSTS AND BIOLOGY 


ALEXEY G. ZINOVJEV AND DAVID R. SMITH 


(AGZ) Zoological Institute, Russian Academy of Sciences, St. Petersburg 199034, Rus- 
sia (e-mail: zag@zisp.spb.su); (DRS) Systematic Entomology Laboratory, PSI, Agricul- 
tural Research Service, U.S. Department of Agriculture, % National Museum of Natural 
History, MRC-168, Washington, DC 20560-0168, U.S.A. (e-mail: drsmith@sel.barc. 
usda.gov) 


Abstract.—The 15 species of sawflies described by Benjamin D. Walsh in 1866 are 
identified. Lectotypes are designated for six species: Euura salicisovum, Euura salicis- 
nodus, Nematus salicispomum, Nematus salicisdesmodioides, Nematus salicispisum, and 
Nematus quercicola. Neotypes are designated for four species: Pristiphora grossulariae, 
Euura perturbans, Nematus inquilinus, and Nematus hospes. Most species described by 
Walsh are the gall-forming Nematinae on willows. Host plants, types of galls, and bio- 


logical notes are presented where known. 


Key Words: 
Walsh 


Benjamin D. Walsh described 15 species 
of sawflies, all of the subfamily Nematinae 
of the Tenihredinidae: Pristiphora grossu- 
lariae (Walsh 1866a) from Davenport, 
Iowa, and 14 species from near Rock Is- 
land, Illinois, in his paper treating insects 
associated with galls on willows (Walsh 
1866b). Three of the latter species, Nematus 
mendicus, N. fur, and Pristiphora_ syco- 
phanta, have free-feeding larvae which 
were only incidental in galls using them as 
a pupation site. All other species described 
in that paper are gall formers. The interpre- 
tation and fixation of the identity of the 
Walsh species by designation of types are 
especially important in the study of North 
American gall-forming sawflies because 
they are among the oldest names available. 
Walsh was the first in North America to de- 
scribe galls and present biological infor- 
mation. 

Walsh’s collection was lost in the Chi- 


Nematus, Pristiphora, Euura, Eupontania, sawflies, galls, willow, Salix, 


cago fire of 1871 (Mallis 1971). However, 
one specimen of Pontania, labeled as the 
type of ‘““Nematus s-pomum Walsh” is in 
the Museum of Comparative Zoology at 
Harvard University, and more Walsh ma- 
terial is in the Academy of Natural Sciences 
of Philadelphia. None of these specimens 
have ever been treated as syntypes and none 
bear Walsh’s original identification label but 
the specimens deposited in Philadelphia fit 
the descriptions and labeling of the speci- 
mens collected and described by Walsh. It 
is improbable that this material had been 
collected by anybody else. Walsh wrote that 
he had sent a part of his material to E. Nor- 
ton (at Philadelphia): “‘I sent a normal fe- 
male of E. perturbans, and a female of E. 
s. ovum to Mr. Norton, along with many 
male and female varieties of Nematus s. po- 
mum, each specimen numbered, but none of 
them named’? (Walsh 1866b: 254). All 
specimens are glued on triangular or pen- 


VOLUME 102, NUMBER 4 


tagonal (in case of Nematus salicispomum) 
cardboard points and each bear a small 
square label with printed letters “‘Ill.”’ (rare 
in handwriting). None of them have iden- 
tification labels written by Walsh himself. 
The labeling varies and identification labels 
(if any) were added by Norton or, for Pon- 
tania species, also by Marlatt. Most speci- 
mens deposited in Philadelphia have small 
squares with numbers, and we assume that 
these are the specimens sent by Walsh to 
Norton. 

In this paper we try to clarify the prob- 
lems concerning sawfly species described 
by Walsh. Lectotypes are selected for six 
species: Euura salicisovum, E. salicisnodus, 
Nematus salicispomum, N. salicisdesmo- 
dioides, N. salicispisum, and N. quercicola; 
a neotype is selected for Pristiphora gros- 
sulariae, and lectotypes of E. salicisovum, 
N. salicispomum, and N. salicisdesmodioi- 
des are designated also as neotypes of E. 
perturbans, N. hospes, and N. inquilinus 
Walsh, respectively. Type material is still 
absent for Euura salicisgemma, E. salici- 
sovulum, N. mendicus, N. fur, and Pristi- 
phora sycophanta. 

Of the 14 names proposed by Walsh, sev- 
en of them can be considered valid: Euura 
salicisovum (= E. perturbans), E. saliciso- 
vulum, E. salicisnodus, Eupontania salicis- 
pomum (= E. hospes), E. salicisdesmodioi- 
des (= N. inquilinus), E. salicispisum (= N. 
quercicola), and Pristiphora sycophanta. 
Pristiphora grossulariae, N. mendicus, and 
N. fur are currently treated as synonyms of 
Pristiphora rufipes Serville, 1823, Nematus 
oligospilus Foerster, and Amauronematus 
histrio Serville, 1823, respectively. How- 
ever, placement of the latter two needs con- 
firmation. 

Most of the gall-making sawflies (and 
midges) described by Walsh have abbrevi- 
ated names. He named many of his species 
like “‘Nematus s. pomum’” or “Euura s. no- 
dus.” Some authors considered them as un- 
available trinomens or used different spell- 
ings for the same species (e.g., Marlatt 
1896, Rohwer 1909, Ross 1951, Smith 


975 


1979). For example, one of the most com- 
mon species creating apple-like galls in 
eastern North America has been placed ei- 
ther in Nematus or Pontania under the 
names *“‘s. pomum Walsh,” “‘salicis-pomum 
Walsh,” ‘“‘pomum Walsh,” ‘‘s-pomum 
Walsh,” or “hospes Walsh (= s. pomum 
Walsh).”’ 

Undoubtedly, “‘s.”” is only an abbrevia- 
tion Walsh used. For the first species in any 
genus, either of midges or sawflies, Walsh 
clearly showed that ‘‘s.”” means “‘salicis. 
For instance: ‘“‘Gall Salicis gemma, n. sp.” 
and then “Euura s. gemma, n. sp.”’ (Walsh 
1866b: 250, No. 16). In fact, these species 
were described as trinomens like ‘“‘Euura 
salicis gemma.’ According to the Interna- 
tional Code of Zoological Nomenclature 
(1985), these are valid names and we 
should accept them, expanding the abbre- 
viation and fusing “‘salicis’”’ with the third 
word. Within sawflies, these names are: Eu- 
ura salicisgemma, E. salicisovum, E. sali- 
cisovulum, E. salicisnodus, Nematus sali- 
cispomum, N. salicisdesmodioides, and N. 
salicispisum. 

Acronyms used for museums are: USNM 
= National Museum of Natural History, 
Smithsonian Institution, Washington, DC; 
MCZ = Museum of Comparative Zoology, 
Harvard University, Cambridge, MA; 
ANSP = Academy of Natural Sciences of 
Philadelphia, PA. Species headings are the 
Walsh species in their original combina- 
tions. 


9 


SPECIES DESCRIBED BY WALSH 


Pristiphora grossulariae Walsh was de- 
scribed from Davenport, Iowa (1866a). All 
other sawfly species described by Walsh 
were published in a paper on insects reared 
from willow galls (1866b) “‘found on sev- 
eral species of Willow in the neighborhood 
of Rock Island, [linois’’ (Walsh 1864). 


Pristiphora grossulariae Walsh 
1866a: 123 
(Figs. 1—2) 
Type material—Described from ‘‘four 
males and forty-nine females” and “forty 


976 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-2. 


3 


larvae of various sizes.’’ One female at 
ANSP probably belongs to the type series 
of this species. It is mounted similar to oth- 
er specimens from Walsh’s collection (Figs. 
1—2), and bears Norton’s identification la- 
bel. We hereby select it as neotype; it is 


Sas 
i don ns 
SET isc A 


Pristiphora grossulariae, neotype. 1, Dorsal view. 2, Lateral view. 


labeled ‘‘Pristiphora grossulariae W. fe- 
male.” 

Valid name.—Pristiphora rufipes (Ser- 
ville 1823) (= Pristiphora grossulariae 
Walsh). Pristiphora rufipes was previously 
attributed to Lepeletier; see Blank and Tae- 


VOLUME 102, NUMBER 4 


ger (1998) for correct authorship of species 
previously attributed to Lepeletier. 

Host.—Ribes spp.; larvae feed on differ- 
ent currant species, but particularly on Ri- 
bes grossularia L. 

Notes.—This is a well-known currant 
pest introduced from Europe. Larvae are 
external leaf feeders. 


Euura salicisgemma Walsh 1866b: 248, 
250 (No. 16) 
(Figs. 3—6) 


Type material.—Described from 34 galls, 
2 larvae, 1 male (reared May 5), and 1 fe- 
male (reared May 2). Specimens possibly 
lost. 

There are no specimens under the name 
“salicisgemma”’ in ANSP, but there are six 
specimens under the name Euura orbitalis 
Norton. All of them are from Illinois, each 
bearing a small square “‘Ill.”” These six 
specimens include one male of Euura, 
which we cannot identify to species, one 
female of Euura salicisovum Walsh, one 
Phyllocolpa sp., and three females of E. or- 
bitalis Norton (= salicisgemma Walsh). 
One of the E. orbitalis specimens is labeled 
we. Am Ent. Soc. Collection’’; 
“TYPE”; “Euura orbitalis’”; “‘Euura orbi- 
talis Norton [Norton’s label]’’; and a label 
“Cannot be Type. Type came from 
“Conn.” SAR” [S.A.Rohwer] (Figs. 3—6). 
This specimen cannot be treated as the type 
of Norton’s Euura orbitalis because Norton 
described it from N.Y. (Brooklyn) and 
Conn., and we cannot treat it as a type of 
Walsh’s species. We were unable to find the 
type of Euura orbitalis Norton elsewhere. 
We doubt if any of these specimens belong 
to Walsh’s material because they are pinned 
and all other Walsh specimens that we have 
seen are glued to points. 

It would be desirable to select neotypes 
(preferably from reared material) for Euura 
orbitalis Norton and Euura_ salicisgemma 
Walsh, to fix the usage of these names, but 
we do not have reared material at present. 

Valid name.—Euura orbitalis Norton 


977 


1862 (= Euura salicisgemma Walsh, syn- 
onymized by Norton 1867). 

Host.—According to Walsh (1866b: 248, 
250), this species creates bud galls on Salix 
humilis Marsh. 

Notes.—This species is related to the 
Holarctic Euura mucronata (Hartig), but E. 
orbitalis is much paler colored (“‘head pale 
luteous’’) than any of the European species 
of Euura. Both species belong to the sub- 
genus Gemmura E. L. Smith (1968), which 
is characterized by a short ovipositor and 
the habit of making bud galls. Euura orbi- 
talis is separated from E. mucronata and E. 
nigrella Rohwer by a short sawsheath in 
dorsal view strongly narrowed at center 
with very short medial flange (Fig. 4). 

We studied specimens from Canada 
(Quebec and Ontario) and U.S.A. (Illinois, 
Missouri, New Hampshire, and New York). 


Euura salicisovum Walsh 1866b: 248, 
251-252 (No. 17) 
(Figs. 7-10) 


Type material.—Described from numer- 
ous galls, 7 larvae, 10 males, and 5 females, 
bred 16—27. 

The lectotype female, here designated, is 
glued on a whitish triangle and labeled 
“Til.”?; “18°. Two paralectotype males are 
labeled 19”; ‘‘Euura s. ovum male IIl. 
Walsh” [on the underside ‘‘gall maker’’]. 
Another paralectotype male is labeled only 
“TH.” (without a number). Deposited in 
ANSP. 

The female lectotype fits Walsh’s de- 
scription. It is in rather good condition, but 
the flagellum of both antennae, the entire 
left hindtarsus, the apical parts of the right 
hindtarsus, the apex of the left midtarsus, 
and the apex of the left forewing are miss- 
ing (Fig. 7). The lower part of the thorax is 
not visible. The identification labels at- 
tached to the male paralectotypes are in 
Norton’s handwriting. There is little doubt 
that the selected lectotype belongs to the 
type series of Euura salicisovum. The only 
other possibility is that it might be a syn- 
type of Euura perturbans Walsh (a syno- 


978 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 3-5. Euura orbitalis, specimen labeled “‘Ill.”’ 3, Dorsal view. 4, Abdomen, dorsal view. 5, Head, dorsal 
view. 


VOLUME 102, NUMBER 4 


Fig. 6. 


Euura orbitalis, head, front view. 


nym of E. salicisovum), but, according to 
Walsh, E. perturbans should be generally 
darker than this female. 

Valid name.—Euura salicisovum Walsh. 

Host.—According to Walsh on “S. cor- 
data.”’ Its currently accepted name is Salix 
eriocephala Michaux (= cordata Muhlen- 
berg not Michaux) (Argus 1997). The galls 
are ‘““‘monothalomous, spongy, growing 
from the side of the twig.’’ Galls obviously 
belonging to this species were common on 
S. eriocephala around Ithaca, NY, in 1997 
(Fig. 10). 

Notes.—The species is pale colored and 
usually larger than Euura orbitalis. It is 
characterized by its longer ovipositor, saw- 
sheath in dorsal view with a more or less 
gradually attenuate medial flange, and the 
entire sawsheath triangular in outline with 
the apical hairs distinctly bent apically (com- 
pare Figs. 4, 9). In both sexes, it is separated 
from E. salicisnodus (together with most of 
other Euura species) by its distinctly con- 
vex, glabrous and strongly shining inner or- 
bits (Fig. 8) and, in dorsal view, the distinct 
inner orbit margins above the eye. 

For further notes, see also Euura salici- 
sovulum Walsh. 


Euura perturbans Walsh 1866b: 254 


Type material.—Described from two 
males and five females bred from cecido- 


O72 


mylidous galls (“‘S. strobiloides, O. S.; “‘S. 
batatus Walsh,” S. rhodoides Walsh (galls 
of the preceding years); and one from a bud 
gall on “Vitis cordifolia.” 

We did not find appropriate specimens in 
ANSP to select a lectotype; however, to fix 
the usage of this name, we hereby select the 
lectotype of Euura salicisovum as neotype 
for E. perturbans. See E. salicisovum for 
labels on the specimen and Figs. 7-9. 

Valid name.—Euura salicisovum Walsh 
(= Euura perturbans Walsh). 

Notes.—Euura perturbans was synony- 
mized by Marlatt (1896). According to 
Walsh, the specimens of E. perturbans 
should be distinguished from E. saliciso- 
vum only by “‘the dorsum of the abdomen 
varying from honey-yellow, including the 
lateral plates, through obfuscated [sic], to 
deep black with the lateral plates also 
black.”’ We consider this a color variation 
and accept the previously proposed synon- 
ymy under E. salicisovum. 


Euura salicisovulum Walsh 1866b: 248, 
253 (No. 18) 


Type material.—Described from 30 galls 
on Salix humilis and 6 larvae. Probably all 
lost. 

Valid name.—Euura 
Walsh. 

Notes.—The galls are the same as those 
of Euura salicisovum; the larvae, according 
to Walsh, are distinguished from those mak- 
ing galls on Salix eriocephala being “all 
decidedly pale greenish,” while those of E. 
salicisovum were “all decidedly yellow- 
ish.”” No galls were found by Walsh on Sa- 
lix discolor which is closely related to S. 
humilis. 

The taxonomic status of this species and 
its host plant specificity are not quite clear. 
Galls of this type occur not only on Salix 
eriocephala and S. humilis (including var. 
tristis), as stated by Walsh, but also on S. 
discolor and S. petiolaris. All such galls 
seem to be associated with a bud, which is 
however left unmodified, the swelling al- 
ways situated just below the bud. 

Specimens that seem to be conspecific 


salicisovulum 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 7-9. Euura salicisovum, lectotype. 7, Dorsal view. 8, Head, front view; arrow indicates inner orbit. 9, 
Apex of abdomen, dorsal view. 


with E. salicisovum are known from Illinois Euura salicisnodus Walsh 1866b: 248, 
and Virginia, but all of them are either cap- 253-254 (No. 19) 

tured or reared from undetermined willows. (Figs. 11-12) 

We are not sure if they belong to E. sali- 

cisovum or to the complex of the sibling Type material.—Described from galls on 
species involving E. saliciovulum. Thus, we ‘‘31 affected twigs,’ undetermined number 
cannot designate a neotype yet. of larvae, and two males. 


VOLUME 102, NUMBER 4 


Fig. 10. Galls of Euura salicisovum on stems of Salix eriocephala, from near Ithaca, NY. 


The lectotype male, here designated is at- 
tached to a whitish elongated triangle and 
labeled “Ill” [in handwriting, on the un- 
derside ““‘B D W #3’’]; “‘Euura salicis nodus 
Walsh.”’ (Fig. 3). Deposited in ANSP. It is 
in rather good condition with only the apex 
of the left flagellum and the left hindtarsus 
missing. 

Valid name.—Euura salicisnodus Walsh. 

Host.—According to Walsh “Salix lon- 
gifolia.”’ Its currently used name is either 
Salix interior Rowlee (= longifolia Muhl- 
enberg) or Salix exigua ssp. interior (Row- 
lee) Cronquist (see Argus 1997). The galls 
are “‘a mere enlargement of the twig, po- 
lythalamous, pitchy inside, with its cells all 
internal” (Fig. 12). 

Notes.—The male lectotype fits the de- 
scription well. It is separated from other 
males of Euura described by Walsh, by 
“black spot enclosing the ocelli is larger, 
and is confluent with the eye for its entire 
length, leaving no orbit between them.”’ 

This species is probably related to E. atra 


Jurine, a species introduced from Europe 
together with its host plants Salix alba L. 
and S. fragilis L. and their hybrids. Both 
species are characterized by the rather flat 
inner orbits and by the narrow sawsheath 
gradually tapering to its apex in dorsal 
view. The longest lateral hairs are distrib- 
uted throughout the entire apical and ven- 
tral part of the sheath (in lateral view) and 
a glabrous medial flange is lacking. Euura 
salicisnodus is separated from E. atra by its 
extensive paler coloration and the galls 
which form an enlargement of the twig. Eu- 
ura atra does not produce enlargements of 
the twig; its galls are practically unnotice- 
able. 

Examined specimens are from Illinois, 
New York, Michigan (?), Ohio, and Canada 
(London, Ontario). Two species described 
from the western United States, Euura mac- 
gillivrayi Rohwer and Euura exigua E. L. 
Smith, are associated with the same willow 
species (or very closely related ones) and 


982 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 11-12. Euura salicisnodis, \ectotype. 11, Dorsal view. 12, Gall on stems of Salix interior, from near 
Ithaca, NY. 


VOLUME 102, NUMBER 4 


they might prove to be only geographical 
forms of E. salicisnodus. 


Nematus salicispomum Walsh 1866b: 248, 
255-256 (No. 20) 
(Figs. 13-16) 


Type material.—Described from very nu- 
merous specimens of galls, an undeter- 
mined amount of larvae, 26 males and 46 
females “‘bred April 16—25 and one female 
bred many years ago.” 

The lectotype female (Figs. 13—16), here 
designated, is on a pentagonal piece of 
brown cardboard, labeled ‘“‘Ill’’; ‘‘12’’; 
“Return to Am. Ent. Soc.”’; “‘Pontania po- 
mum Walsh’’; “‘N. s. pomum Walsh.” De- 
posited in ANSP. 

In the ANSP, there are 11 paralectotypes, 
6 females and 5 males, all mounted on sim- 
ilar pentagonal cardboard points: 2 females 
and 3 males are without a number, and 1 
female is labeled “Hl:; 1527; “Return to 
Am. Ent. Soc.” The heads of one male and 
of one female are missing, otherwise they 
are in good condition. One female labeled 
eile EYPE 14013": “Cresson”; “Ne- 
matus s-pomum Walsh”’; ““MCZ // Museum 
of Comparative Zoology” is deposited in 
the MCZ. Two females and 2 males, all on 
similar cardboard points and similarly la- 
beled “‘Ill.’’; “‘Return to Am. Ent. Soc.” 
with two of them bearing C. L. Marlatt’s 
identification labels ‘‘Pontania pomum 
Walsh” are deposited in the USNM. 

Valid name.—Eupontania salicispomum 
(Walsh), new combination. See Zinovjev 
(1993) for the definition of Eupontania. 

Host.—The galls are on Salix eriocepha- 
la Michaux (= S. cordata). A subsidiary 
host plant might be Salix discolor Muhl- 
enberg. Walsh wrote about it as follows: 
“On S. cordata, (and very rarely on Salix 
discolor)”; he found galls of Nematus sal- 
icispisum and galls “‘so identical in appear- 
ance with S. pomum’’ on the same bushes.” 
“In both the above two cases a few discolor 
bushes were growing in the midst of very 
large numbers of S. cordata, the species on 
which S. pomum is normally found.’’ These 


983 


data on its host plant specificity seem to be 
reliable, but confirmation by rearing adults 
is needed. 

The galls are roundish (apple-like), tran- 
sected by the leaf blade with a large part of 
the gall visible from the upper side of the 
leaf, but the larger part of the gall is situated 
below leaf surface (illustrated by Zinovjev 
and Smith 1999: 361, fig. 2). 

Notes.—In this species, the hind tibial 
spur is shorter than the apical tibial breadth, 
the frontal area lacks hairs on its anterior 
part, the inner orbits are glabrous and shiny 
(Fig. 15) with hairs only along the eye mar- 
gin, and the sawsheath in dorsal view is a 
narrow triangle with rounded margins (Fig. 
16). The frontal wall of the frontal area is 
usually considerably protruding (e.g., in 
reared specimens from New York), but in 
specimens of the type series, the whole up- 
per head in lateral view is rather flat. 


Nematus salicisdesmodioides Walsh 
1866b: 248, 257-258 (No. 20) 
(Fig. 17) 


Type material—Described from an un- 
determined number of galls, “three or 
four” larvae, 2 males and 8 females, bred 
April 2-15. 

The lectotype female (Fig. 17), here des- 
ignated, is attached to an elongated triangle 
and labeled ‘“‘Ill.’’; “13’’; ““Return to Am. 
Ent. Soc.”’; ““‘Pontania desmodioides female 
Wish.” [identification label of Marlatt]; “‘N. 
desmodioides Walsh [identification label of 
Norton]. Deposited in ANSP. 

A paralectotype male is deposited in 
ANSP. It is mounted similar to the female, 
and it is labeled the same (with number 13), 
but without an identification label. 

Valid name.—Eupontania salicisdesmo- 
dioides (Walsh), new combination. 

Host.—Salix humilis Marshall. 

Notes.—The galls are ‘“‘semicircular in 
outline, sessile’? (transected by the leaf 
blade and equally developed on both sides 
of the leaf). 

The female can be separated from other 


984 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 13-16. 


Eupontania by the following combination 
of characters: sawsheath in dorsal view as 
a long triangle with an acute apex and 
straight sides and with hairs directed more 
posteriorly than laterally (with an angle of 
less than 90°); upper part of head in lateral 


Eupontania salicispomum, \ectotype. 13, Dorsal view. 14, Head, dorsal view. 15, Head, front 
view; arrow indicated inner orbit. 16, Apex of abdomen, dorsal view. 


view densely covered by hairs thoughout, 
hind tibial spurs shorter than apical tibial 
breadth; and inner orbits with dense hairs. 

An additional female was examined from 
Massachusetts (deposited in ANSP) [seen 
by Marlatt because of his label]. 


VOLUME 102, NUMBER 4 


985 


Fig. 17. 


Nematus salicispisum Walsh 1866b: 248, 
258-260 (No. 21bis) 
(Figs. 18—20) 


Type material.—Described from many 
galls, larvae, two males, and three females. 

The lectotype female, here designated 
(Figs. 18-20), is attached to a long trian- 
gular cardboard point, labeled “‘Ill.’’; “14”; 
“Return to Am. Ent. Soc.’’ Two paralec- 
totypes males are similarly labeled as the 
lectotype; one is labeled number 14 as is 
the lectotype, and the other is labeled num- 
ber 15. 

Valid name.—Eupontania salicispisum 
(Walsh), new combination. 

Host.—Salix discolor: The gall is illus- 
trated by Zinovjev and Smith (1999: fig. 1). 
They are small, rounded, and attached to 
the sideveins, with a very small scar visible 
from the upperside and concave below the 
leaf surface. 

Notes.—In the key to Palearctic species 
(Zinovjev 1993) this species runs to the vi- 


Eupontania salicisdesmodiodes, \ectotype, dorsal view. 


minalis group (length of inner hind tibial 
spur subequal to maximum width of hind 
tibial apex and rather thin). It is character- 
ized also by its small size; short ovipositor 
(Fig. 20); sawsheath in dorsal view as a 
short triangle with rounded sides (Fig. 19) 
with the longest hairs directed more later- 
ally than posteriorly and more strongly bent 
near their apices; upper part of head in lat- 
eral view with dense erect hairs, but front 
wall of frontal area glabrous; antennal hol- 
low glabrous and shining; and inner orbits 
with hairs well developed. 


Nematus quercicola Walsh 1866b: 260 
(Figs. 21—24) 


Type material.—Described from 2 males, 
7 females reared from ‘‘an undescribed, 
cabbage-like, polythalamous, Cecidomyi- 
idous gall on the White Oak ...” 

The lectotype (Figs. 21—24) is a female 
labeled “‘IIl’’; ““N. quercicola Walsh” [Nor- 
ton’s label]; ‘“‘Return to Am. Ent. Soc.”’; 


986 


Figs. 18—20. 
Apex of abdomen, lateral view. 


‘‘Pontania pisum Wlsh female” [Marlatt’s 
identification label]. Deposited in ANSP. 
Valid name.—Eupontania salicispisum 
(Walsh) (= Nematus quercicola Walsh). 
Notes.—Nematus quercicola was treated 
as a synonym of the preceding species by 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Eupontania salicispisum, lectotype. 18, Dorsal view. 19, Apex of abdomen, dorsal view. 20, 


Marlatt (1896), or by others (e.g., Smith 
1979) as a nomen nudum because the only 
character mentioned by Walsh to distin- 
guish it from the preceding species was that 
‘tall N. s. pisum went underground to pup- 


> 


ize’? and ‘‘N. quercicola pupized in the 


VOLUME 102, NUMBER 4 987 


22 


Figs. 21-24. Nematus quercicola, lectotype. 21, Dorsal view. 22, Apex of abdomen, dorsal view. 23, Head 
and thorax, lateral view. 24, Abdomen, lateral view. 


988 


gall.”” A biological feature such as this, by 
formal reasons, could be treated as a “‘char- 
acter,” and we should accept this name as 
an available one. According to Walsh, this 
species ‘“‘cannot be distinguished from the 
gallmaking N. s. pisum male, female.” 


Nematus inquilinus Walsh 1866b: 
260-261 


Type material.—Described from one 
male, three females, bred from the cecido- 
myiidous gall “‘S. rhodoides Walsh. No ap- 
propriate material was found in ANSP. The 
neotype female, here designated is the lec- 
totype of Nematus salicisdesmodiodes 
Walsh (Fig. 17). 

Valid name.—Eupontania salicisdesmo- 
diodes (Walsh) (= Nematus inquilinus 
Walsh). 

Notes.—This species was placed in syn- 
onymy with Nematus salicisdesmodioides 
Walsh (Marlatt 1896) and was listed as 
‘“‘unplaced”’ by Smith (1979). The type ma- 
terial was reared from galls on Salix hu- 
milis, the host plant of Eupontania salicis- 
desmodioides. To fix the usage of this name 
we designate the lectotype of Nematus sal- 
icisdesmodioides as neotype of Nematus in- 
quilinus Walsh. 


Nematus hospes Walsh 1866b: 261 


Type material.—Described from one 
male and two females from the gall of s. 
strobiloides Osten-Sacken. No type mate- 
rial was located, but there is no strict evi- 
dence that absolutely all specimens under 
the name WN. salicispomum belong to the 
original type series. To fix the usage of this 
name we select the lectotype of N. salicis- 
pomum Walsh (Figs. 13—16) as the neotype 
of N. hospes Walsh. 

Valid name.—Eupontania salicispomum 
(Walsh) (= Nematus hospes Walsh). 

Notes.—According to Walsh ‘‘absolutely 
undistinguishable from the normal type of 
the gall-making N. s. pomum.” Marlatt 
(1896) synonymized N. hospes, and these 
species have always been treated as con- 
specific. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Nematus mendicus Walsh 1866: 261 


Type material.—Described from one 
male, three females: ‘“‘one female bred May 
2 from the Tenthredinous gall S. pomum n. 
sp. of the preceding year’s growth, and an- 
other female, August 5, from the Cecido- 
myidous gall S. brassicoides Walsh of the 
same year growth; the other female and the 
male captured at large.’’ No type material 
was located. 

Valid name.—Nematus oligospilus Foers- 
ter 1854 (= Nematus mendicus Walsh). 

Host.—Willows, Salix spp. 

Notes.—Benson (1962) synonymized 
Nematus mendicus, thus treating N. oligos- 
pilus as a Holarctic species. It is in a com- 
plex of green Nematus species with external 
leaf-feeding larvae. Further studies may re- 
veal several species in this complex. Cur- 
rently, we accept Benson’s synonymy. 


Nematus fur Walsh 1866: 263 


Type material.—Described from a single 
male ‘“‘bred March 29 from an old bored 
subpeduncled spherical gall, .57 inch in di- 
ameter, made by Cecidomyia s. batataus 
Walsh on S. humilis.” The type was not lo- 
cated. 

Valid name.—Currently, Nematus fur is 
treated as a synonym of Amauronematus 
histrio Serville 1823. However, its taxo- 
nomic position is not quite clear. It is pos- 
sibly a valid species of Amauronematus. 

Notes.—This species was treated by 
Ross (1951) as a synonym of Amaurone- 
matus luteotergum Norton 1861, and by 
Smith (1979) as synonym of A. histrio Le- 
peletier 1823 (= luteotergum Norton). 
However, these treatments do not fit exactly 
the description of N. fur. According to 
Walsh, Nematus fur has “‘black legs” and 
‘“‘wings subhyaline, slightly tinged with fu- 
liginous.”’ This dark coloration of the wings 
and legs is rare in the Nematinae, and we 
could not find appropriate specimens in col- 
lections that fit the description of N. fur. 
Otherwise, according to its morphology 
(particularly microsculpture) and its ability 


VOLUME 102, NUMBER 4 


to bore for pupation into plant tissue, it al- 
most certainly belongs to Amauronematus. 


Pristiphora sycophanta Walsh 1866b: 263 


Type material.—Described from a single 
male “‘bred August 9 from a cocoon found, 
July 27, inside the Cecidomyidous gall S. 
brassicoides Walsh of the same year 
growth.” The type was not found. 

Valid name.—Pristiphora sycophanta 
Walsh. 

Host plants.—This is a common Pristi- 
phora on willows. The cecidomyidous gall 
from which the type was reared was on Sa- 
lix interior. 

Notes.—Numerous specimens were tak- 
en in Clarke Co., VA, where the predomi- 
nant willows around a pond were Salix ni- 
gra Marsh. Malaise trap collections at this 
locality included P. sycophanta specimens 
throughout the spring and summer, indicat- 
ing that there are several generation a year. 
This supports our interpretation of P. sy- 
cophanta, which Walsh stated was reared 
from the same years growth. 


ACKNOWLEDGMENTS 


We thank D. Azuma (Academy of Nat- 
ural Sciences of Philadelphia, PA) and S. 
Cover (Museum of Comparative Zoology, 
Harvard University, Cambridge, MA) for 
allowing study of important material in 
their collection, and I. M. Kerzhner (Zoo- 
logical Institute, Russian Academy of Sci- 
ences, St. Petersburg) for advice on usage 
of the Walsh names. This study was sup- 
ported in part by Smithsonian Instituion 
Short-Term Visitors Grants, 1997 and1999, 
to A.G. Zinovjev. We thank Terry Nuhn 
(Systematic Entomology Laboratory, USDA) 
for taking the photographs and Cathy An- 
derson (Systematic Entomology Laborato- 
ry, USDA) for arranging the plates. We ap- 
preciate the comments of the following re- 
viewers: Henri Goulet (Agriculture and 
Agri-Food Canada, Ottawa) and T, J. Henry 
(Systematic Entomology Laboratory, 
USDA). 


989 


LITERATURE CITED 


Argus, G. W. 1997. Infrageneric classification of Salix 
(Salicaceae) in the New World. Systematic Botany 
Monographs 52: 1-121. 

Benson, R. B. 1962. Holarctic sawflies (Hymenoptera: 
Symphyta). Bulletin of the British Museum (Nat- 
ural History) Entomology 12: 381—409. 

Blank S. M. and A. Taeger. 1998. Comments on the 
taxonomy of Symphyta (Hymenoptera) (Prelimi- 
nary studies for a catalogue of Symphyta, part. 4), 
pp.141—174. In Taeger, A. and S. M. Blank, eds., 
Pflanzenwespen Deutschlands (Hymenoptera, 
Symphyta). Kommentierte Bestandsaufnahme. 
Verlag Goecke & Evers, Keltern, 368 pp., 4/5 ta. 

International Code of Zoological Nomencalture. 1985. 
University of California Press, Berkeley and Los 
Angeles, 338 pp. 

Marlatt, C. L. 1896. Revision of the Nematinae of 
North America, a subfamily of leaf-feeding Hy- 
menoptera of the family Tenthredinidae. United 
States Department of Agriculture, Division of En- 
tomology, Technical Series No. 3, 135 pp. 

Mallis, A. 1971. American Entomologists. Rutgers 
University Press, New Brunswick, New Jersey, 
549 pp. 

Norton, E. 1862. Notice of several new species of Ten- 
thredinidae. Proceedings of the Entomological So- 
ciety of Philadelphia 1: 143-144. 

. 1867. Catalogue of the described Tenthredi- 
nidae and Uroceridae of North America. Trans- 
actions of the American Entomological Society 1: 
193-280. 

Rohwer, S. A. 1909. The sawfly genus Cryptocampus 
in boreal North America. Journal of the New York 
Entomological Society 17: 7—25. 

Ross, H. H. 1951. Symphyta, pp. 4-89. Jn Muesebeck, 
C. E W., K. V. Krombein, and H. K. Townes, eds., 
Hymenoptera of America North of Mexico, Syn- 
optic Catalog. United States Department of Agri- 
culture, Agriculture Monograph No. 2, 1420 pp. 

Smith, D. R. 1979. Symphyta, pp. 3-137. In Krom- 
bein, D. R., P. D. Hurd, Jr, D. R. Smith, and B. 
D. Burks, eds., Catalog of Hymenoptera in Amer- 
ica North of Mexico, Vol. 1. Smithsonian Insti- 
tution Press, Washington, D.C. 

Smith, E. L. 1968. Biosystematics and morphology of 
Symphyta. I. Stem-galling Euura of the California 
region, and a new female genitalic nomenclature. 
Annals of the Entomological Society of America 
61: 1389-1407. 

Walsh, B. D. 1864. On the insects, coleopterous, hy- 
menopterous and dipterous, inhabiting the galls of 
certain species of willow. Part 1st.—Diptera. Pro- 
ceedings of the Entomological Society of Phila- 
delphia 3: 543-644. 

1866a. Imported insects;—The Gooseberry 

sawfly. The Practical Entomologist. A Monthly 

Bulletin 1(12): 117-125. 


990 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


. 1866b. On the insects, coleopterous, hyme- viminalis-group (Hymenoptera: Tenthredinidae). 
nopterous and dipterous, inhabiting the galls of Zoosystematica Rossica 2: 145—154. 
certain species of willow.—Part 2d and last. Pro- _Zinovjev, A. G. and D. R. Smith. 1999. Types and 
ceedings of the Entomological Society of Phila- biological notes of the eastern North American 
delphia 6: 223-288. sawflies of Pontania Costa and Phyllocolpa Ben- 
Zinovjev, A. G. 1993. Subgenera and Palaearctic spe- son (Hymenoptera: Tenthredinidae) described by 
cies groups of the genus Pontania, with notes on Marlatt, Dyar, and Rohwer. Proceedings of the En- 


the taxonomy of some European species of the tomological Society of Washington 101: 359-371. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 991-1002 


A NEW SPECIES OF THE SUBGENUS SABETHOIDES OF SABETHES 
(DIPTERA: CULICIDAE) FROM VENEZUELA AND BRAZIL 


Dusty A. MOosgEs, THERESA M. HOWARD, AND RALPH E. HARBACH 


(DAM) Department of Biology, Cumberland College, 7000 College Station Drive, Wil- 
liamsburg, KY 40769, U.S.A.; (TMH, REH) Department of Entomology, The Natural 
History Museum, Cromwell Road, London SW7 5BD, U.K. (e-mail: t-howard@nhm.ac. 


uk; r.harbach@nhm.ac.uk) 


Abstract.—The adult male, pupa and larva of Sabethes (Sabethoides) conditus, n. sp., 
are described from localities in western Brazil and northern Venezuela. The species is 
distinguished from Sabethes chloropterus (von Humboldt), which it closely resembles in 


all life stages. 


Key Words: Diptera, Culicidae, Sabethes, 


Venezuela 


Mosquitoes of subgenus Sabethoides 
Theobald of genus Sabethes Robineau-Des- 
voidy are some of the more common insects 
in Neotropical forests, yet they are poorly 
known and rarely studied. No significant 
taxonomic research has been done on the 
subgenus since Lane (1953). This is sur- 
prising since one of the species, Sa. chlo- 
ropterus (von Humboldt), has repeatedly 
been found infected with St. Louis enceph- 
alitis virus (Galindo et al. 1959) and is 
known to transmit yellow fever virus to hu- 
mans (Galindo et al. 1956). In addition to 
Sa. chloropterus, the subgenus includes Sa. 
glaucodaemon (Dyar and Shannon), Sa. tri- 
dentatus Cerqueira, and at least 2 undescri- 
bed species. With the addition of the new 
species described in this paper, Sabethoides 
now includes 4 formally recognized spe- 
cies. 

The new species described here very 
closely resembles Sa. chloropterus in all 
life stages, and undoubtedly has been con- 
fused as this species in the past. For this 


Sabethoides, new species, mosquito, Brazil, 


reason, future study may indicate that it is 
also a vector of arboviruses. 


MATERIALS AND METHODS 


This study is based on specimens bor- 
rowed from the National Museum of Nat- 
ural History, Smithsonian Institution, 
Washington, DC. Observations of the adults 
were made under simulated natural light. 
Larval and pupal chaetotaxy were studied 
using a combination of bright field and dif- 
ferential interference contrast microscopy. 
Measurements and counts were taken from 
all specimens of the type series where the 
structures in question were present. Num- 
bers in parentheses represent modes of the 
reported ranges unless indicated otherwise. 
The form of presentation, descriptive ter- 
minology and abbreviations used in the spe- 
cies description follow Harbach and Knight 
(1980, 1982) and recent papers published as 
part of an ongoing revision of the genus 
Sabethes (Harbach and Peyton 1991; Har- 


992 


A B C D 


Fig. 1. Tibia and tarsus of midleg of Sabethes (Sa- 
bethoides) conditus (A, B) and Sa. chloropterus (C, 
D). A, C, Anterior; B, C, posterior. Scale = 2.0 mm. 


bach 1991la, 1994, 1995a, b; Harbach and 
Petersen 1992; Hall et al. 1999). 


Sabethes (Sabethoides) conditus, Moses, 
Howard, and Harbach, new species 
(Figs. 1—4) 


Sabethes (Sabethoides) sp 4 of Heinemann 
and Belkin 1978: 373. 


This species exhibits the diagnostic char- 
acteristics of the subgenus noted by Har- 
bach (1991a): midtarsus marked with white 
scaling; prespiracular, upper proepisternal 
and lower mesokatepisternal setae present; 
prealar setae absent; and legs without pad- 
dles. 

Male.—Medium-sized mosquito with 
brilliant metallic-coloured scaling; scales of 
head capsule, thorax and abdomen very 
broad and flat; scales of vertex with differ- 
ent combinations of metallic blue, violet 
and green reflections depending on angle of 
light; scales of postgena, thoracic pleura 
and coxae silvery white; antepronotum 
ranging from dark gold to bright cerulean 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


blue depending on angle of light; scaling of 
postpronotum similar to pleura but with a 
slight golden hue; scutal and scutellar scales 
metallic gold with cerulean reflections; me- 
sopostnotum without scales; proboscis and 
legs predominantly metallic blue and violet, 
proboscis darker; wing entirely dark-scaled 
with blue, gold and violet iridescence when 
viewed from certain angles; abdominal ter- 
ga primarily metallic gold with narrow pale 
basal bands, from dorsal angles golden ar- 
eas appear cerulean to green and basal 
bands mauve to violet, basal bands broader 
laterally and silvery-white in lateral view. 
Head: Eyes joined above and below. Oc- 
ciput with transverse row of short semi- 
erect scales at back of head. Ocular setae 
short, dark, close to margin of eye; 2 long, 
bronzy, approximated interocular setae pre- 
sent. Antenna: Dark; length 1.29—1.36 mm 
(x = 1.34 mm), significantly shorter than 
proboscis; pedicel large, surface silvery pu- 
bescent; flagellum rather strongly verticil- 
late, proximal whorls with 9 or 10 setae, 
longest setae about 0.33 length of antenna. 
Clypeus and frons without setae and scales, 
dark. Proboscis long, slender, straight; 
length 2.00—2.10 mm (* = 2.05 mm); distal 
0.3 gradually expanded laterally, becoming 
twice as broad as proximal part; dark-scaled 
with ventral patch of yellowish white scal- 
ing extending 0.5 to 0.8 from base. Maxil- 
lary palpus short, about 0.08 length of pro- 
boscis; silvery-scaled dorsally, bare ven- 
trally. Thorax: Integument brown. Dorsum 
with dark setae on anterior promontory (10, 
11), antepronotum (11-13), supraalar area 
(10-15), scutellum (3,4 long setae on lat- 
eral lobes; 2 long setae projecting down- 
ward from midlobe) and mesopostnotum 
(4). Pleura with prespiracular (2), upper 
proepisternal (1), lower mesokatepisternal 
(1,2) and upper mesepimeral setae (9-11); 
prespiracular setae dark, others yellow or 
golden. Lower part of proepisternum with- 
out scales, scales on upper part contiguous 
with scales on anteprocoxal membrane; 
scales present on upper portion of postpro- 
coxal membrane; mesopleuron with scales 


VOLUME 102, NUMBER 4 


covering all but lower anterior margin of 
mesokatepisternum, extreme dorsal margin 
of postspiracular area, narrow ventral and 
upper posterior margins of mesepimeron 
and mesomeron; scales absent from meta- 
pleuron, metameron and postmetacoxal 
membrane. Wing: Length 3.2 mm; dorsal 
scales broader and slightly asymmetrical on 
anterior and distal veins; alula with fine pil- 
iform scales on margin distally; calypters 
without setae. Halter: Scabellum without 
scales, integument pale; pedicel and capi- 
tellum dark-scaled. Legs: Coxae and tro- 
chanters with silvery-white scales, trochan- 
ters with some dark scales dorsally at apex; 
femora dark above and golden below; all 
tibiae and fore- and hindtarsi entirely dark- 
scaled; anterior surface of midtarsus (Fig. 
1) white-scaled from middle of tarsomere 2 
to middle of tarsomere 4 (paratype from 
Brazil with white scaling encircling tarso- 
mere 3 and part of 4). Forefemur about 1.2 
length of proboscis, same length as midfe- 
mur, about 1.3 length of hindfemur; hind- 
tibia about as long as hindfemur, hindtar- 
somere 1 about 1.2 length of hindfemur. 
Ungues small, simple, black. Abdomen: 
Coloration as noted above, sternal scales 
distinctly larger and less recumbent than 
tergal scales. Genitalia (Fig. 2): Tergum 
VIif (ventral in position; not illustrated) 
with deep V-shaped emargination at middle 
of posterior margin, posterior border on ei- 
ther side of emargination with 3 or 4 irreg- 
ular rows of long close-set setae, posterior 
half of surface before setae covered with 
recumbent spatulate scales that rather 
abruptly grade into a cluster of much larger 
scales on posterolateral corners, scales of 
cluster about half length of marginal setae. 
Tergum and sternum IX fused laterally, 
forming a complete ring of sclerotization; 
ninth tergal lobes small, not produced, each 
bearing 3 flattened setae with apices bent 
laterad; interlobular bridge moderately 
broad. Gonocoxite stout, width more than 
half length in lateral view, tapered in distal 
third, tergomesal surface membranous, dis- 
tal sternal area covered with scales and 


993 


short setae, bearing 3 long tergomesal setae 
below basal mesal lobe; basal mesal lobe of 
irregular shape, roughly trapezoidal in ven- 
tral (tergal) view, partly covered with small 
slender setae and bearing 2 large setae at 
caudolateral angle. Gonostylus (side view) 
large, about two-thirds length of gonocox- 
ite; stem stout, short, less than half length 
of head; head as illustrated, bearing 5 well 
developed lobes (A, C, E, M, M’), a slight- 
ly produced lobe B and an elongate tergal 
fimbriate process; lobe A, elongate tapered 
process arising tergomesal of base of lobe 
M, bearing several short stout setae at apex; 
lobe E, rather short conical lobe borne mes- 
ally at bases of lobes A and M, with large 
blade-like seta and 2 smaller needle-like se- 
tae at apex; lobe B, lateral minutely spicu- 
late swelling associated with sternomesal 
margin of lobe E, bearing 3 large sternally 
directed setae with distal ends flattened and 
expanded subapically; lobe M, irregular 
partially subdivided median apical lobe 
bearing broadly V-shaped tergal process 
with short striated arms and more proximal 
long slender tergally directed fimbriate pro- 
cess comprised of partially fused and coa- 
lesced filaments with free ends bending to- 
ward base of gonostylus; lobe M’, elongate 
irregular process arising from sternolateral 
margin at base of lobe M, subapical tergo- 
lateral margin with several short thickened 
setae and more proximal small flattened 
sigmoidal seta, sternal margin with laterally 
compressed projection bearing a narrowed 
retrorse apex; lobe C, large mesally slanted 
process arising from sternal margin at base 
of lobe M’, bearing pattern of decumbent 
spine-like spicules on mesal side of apical 
margin and cluster of long sternally direct- 
ed filaments on lateral side at proximal end. 
Aedeagus longer than wide, widest in basal 
half; submedian tergal arms fused to form 
broad median tergal bridge; apical tergal 
arms fused to form narrow apical bridge; 
median sternal plate rather membranous, 
apex not markedly flared, hood-like. Proc- 
tiger (in lateral view) with broad basal 
sclerotization (tergum X); paraproct slender, 


994 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


lateral 
tergal 


lateral 


tergal 


Fig. 2. Male genitalia of Sabethes (Sabethoides) conditus. Aspects as indicated for (A) gonocoxopodite; (B) 


tergum IX; (C) gonostylus; (D) proctiger; (E) aedeagus, with parameres attached. A, B, C, E, M and M’ = 
gonostylar lobes; BML = basal mesal lobe; fp = fimbriate process; tms = tergomesal setae. Scale in mm. 


VOLUME 102, NUMBER 4 


N 


LE 


\ 


Yigg Teilard. 


Fig. 3. Fourth-instar larva of Sabethes (Sabethoides) conditus. A, Head, dorsal (left) and ventral (right) 
aspects of left side. B, Dorsomentum. C, Thorax and abdominal segments I-VI, dorsal (left) and ventral (right) 


aspects of left side. D, Abdominal segments VII-X, left side. E, Comb scale. A = antenna; C = cranium; CS 
= comb scale; Dm = 


dorsomentum; P = prothorax; M = mesothorax; S = siphon; T = metathorax; I-X = 
abdominal segments; 1-15 = setal numbers for specified areas, e.g., seta 5-C. Scales in mm. 


995 


996 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 1. Numbers of branches for setae of fourth-instar larvae of Sabethes (Sabethoides) conditus. 
Head Thorax Abdominal segments 
Seta oc re, ae P M T I il 
0) 1 10-13 — -- a i 
1 1 5,6(5,6) 3—5(4) 3 2,3(3) Dy 
2 — | 1 1 DES (2-3)) 3,4(3) 
3 1 2 l 2,3(2) 1 1 
4 1 6—9(7) i 2-4(2) 2—4(3) 4—6(4) 
5 D, 1,2(1) ] 3—5(3) 3 5,6(6) 
6 1 2, 1 1—3(2) 5,6(6) 5—8(5) 
7 3—5(3) 7,10(10) i 10—12(10) 5-7(5) 46(6) 
8 DES (23) 9-11(10) 5,6(6) 10—12(10) — 1,21) 
9 y 2,3(2) 1,3(1) 7,8(7,8) 1 2 
10 1e2@) 6—8(7) 5,6(5) 1 ] 1 
11 6—-9(9) 1,2(2) 1,2(2) 1,2(1) 6—-8(7) 4 
12 3,4(3) 1 1 1 — ] 
13 1 — 13-16 5,6(6) 6,7(6) 5—7(6) 
14 4,5(5) 2—4(4) 12—-19(19) — — -—— 
15 1—3(1) = = = = = 


apex with 2 or 3 appressed teeth, rather 
large subapical area distinctly sclerotized, 
bearing 6—23 cercal setae. 

Female.—Not definitely associated with 
the male (see Systematics, below); 2 pre- 
sumed females from Brazil resemble the 
males from Venezuela except for the ab- 
sence of a pale patch on the ventral surface 
of the proboscis and in having slightly more 
extensive pale scaling on the anterior sur- 
face of the midtarsus, extending from prox- 
imal 0.2 of tarsomere 2 to apex of 4. 

Egg.—Unknown. 

Larva, fourth instar (Fig. 3).—Exhibiting 
the subgeneric characters noted by Harbach 
1991a; similar to Sa. chloropterus (cf. Fig. 
4 in Harbach 1991b); character and place- 
ment of setae as figured, numbers of 
branches in Table 1. Head: Slightly wider 
than long, widest in posterior half; length 
about 1.0 mm; width about 1.3 mm. Occip- 
ital foramen widely V-shaped with arms ex- 
tending dorsolaterally to point laterad of 
level of seta 9-C, margins heavily tanned, 
ventrocaudal margin with collar-like edge. 
Anterior margin of labiogula weakly den- 
ticulate; hypostomal suture complete, gent- 
ly curved. Dorsomentum (Fig. 3B) short, 
with 6,7 teeth on either side of median 


tooth, median tooth and most lateral tooth 
larger than others. Maxilla, including apical 
tooth, about half length of head, with 8-10 
lateral teeth, first lateral tooth larger than 
the others which become progressively 
smaller. Setae 4,6-C single, simple; 7-C 
usually triple; 9-C inserted at level distinct- 
ly posterior to 10-C; 10-C single or double; 
14-C without thickened branches; 15-C in- 
serted cephalad of 14-C near anterior mar- 
gin of labiogula, long with 1—3(1) branches. 
Antenna: Short, cylindrical; slightly ta- 
pered distally, length about 0.28 mm. Seta 
1-A single, simple, borne dorsally about 0.8 
from base, length about 2 times width of 
antenna at point of insertion. Thorax: In- 
tegument hyaline, smooth. Setae 0,1,8,14-P, 
1,13,14-M and 5,8-T with multiple short 
flexible branches; 11-P,.M,T single or dou- 
ble, 11-R.M with slender flexible branches, 
11-T with stiff thickened branches; 13-T 
with multiple branches, about length of tho- 
rax. Abdomen: Integument hyaline, smooth 
except for rows of minute spicules before 
comb. Seta 1-I,II with thickened branches, 
1-III-V long, double with one branch mark- 
edly shorter; 2-I laterad of seta 1, 2-II-VII 
well mesad and anterior to seta 1; 6,7-I, I 
similar, strongly developed with multiple 


VOLUME 102, NUMBER 4 997 
Table 1. Continued. 
Abdominal segments 
= we) vy hh. vw. = ww wt hoo 
1 1 1 1 1 1 = 
2 2 2 2 223 (2) 5—8(5) 2 
3,4(3,4) 3,4(4) 3)9\G)5) 3,4(3) 23D) 1 3,4(3,4) 
1 ] 1 1 1 6,7(6,7) 2 
] 1 8—11(8) HED En) 1 1 2 
4,5(4,5) 3,4(4) 3,4(3) D2) 2—5(3) 2 — 
1 ] 1 1 10—-14(13) — 
9-12 11-16 11—13(12) 5—8(7) ] a — 
2,3(3) 2 1,2(2) 14-17 19-25 1-S, 1 
23 (2-5) 23) 2 SPX) 2) la-S, 1 
1 1 1,2(1) i 1,2(2) 2a-S, 1 
4,5(4,5) 3,4(3) 2-6 15-21 1 a — 
1 1 | 1 1 oe 
5-7(7) 5—6(5,6) 5,6(5) 29—34(29) 3,4(3) = = 


aciculate branches, 6-III-VI long, single and 
progressively shorter; 5-II-VI and 13-I-V 
strongly developed, stellate, with stiff acic- 
ulate branches; 13-III-V apparently with 
more branches (x = 17) than Sa. chlorop- 
terus (x 12); punctures present on seg- 
ments III-V. Segment VIII: Comb a single 
row of 21—26 long slender spine-like scales 
(Fig. 3E) without fringe of minute spicules, 
scales close-set with flared bases sometimes 
partially joined. Siphon: Relatively long 
and slender, gradually tapered from base to 
apex; moderately tanned, surface more or 
less evenly covered with short rows of mi- 
nute spicules; length about 1.3 mm, width 
at mid-length about 0.1 mm, index about 
13. Pecten of about 30 fine filaments ex- 
tending from below level of insertion of 
seta 1-S to point distal to seta la-S. Seta 1- 
S inserted 0.25 from base of siphon, la-S 
includes 1 or 2, usually 2, and 2a-S includes 
2 or 3, usually 3, single setae; seta 2-S 
stout, slightly sinuous and minutely forked 
at tip (not apparent in Fig. 3D). Segment X: 
Saddle borne on posterodorsal quarter of 
segment, surface more or less evenly cov- 
ered with rows of minute spicules; length 
about 0.2 mm; siphon/saddle index about 


6.5. Setae 1—4-X equally well developed; 
1,3,4-X double, 2-X usually triple (3,4). 
Pupa (Fig. 4).—Exhibiting the subgener- 
ic characters noted by Harbach 1991a; sim- 
ilar to Sa. chloropterus; character and po- 
sitions of setae as illustrated, numbers of 
branches in Table 2. Cephalothorax: Light- 
ly tanned. Seta 1-CT strongly developed, 
double, branches sigmoidally curved with 
hooked tips; 5-CT also well developed, 
double or triple, more often triple. Trumpet: 
Moderately tanned, slightly flattened but lit- 
tle if at all expanded laterally; length 0.32— 
0.39 mm (x 0.35 mm), width at mid- 
length 0.11—0.14 mm, index 2.78—-2.91 (x 
= 2.81); pinna short, length about 0.08 mm. 
Abdomen: Lightly tanned, terga and sterna 
darker anteriorly; length about 3.7 mm. 
Seta 1-I well developed, with about 80 ter- 
minal branches; 6,7-I long, 6-I single, lon- 
ger than 7-I; 2-II-VII lateral to seta 1; 5- 
II, with multiple branches, much shorter 
than 5-IV-VI, 5-IV-VI very long, more than 
twice length of following tergum; seta 4-IV 
short, with multiple branches, generally 
with more branches than in Sa. chloropte- 
rus which usually has 1 or 2; 10-II and 13- 
VI present on both sides in 2 available 


998 


Table 2. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Numbers of branches for setae of pupae of Sabethes (Sabethoides) conditus. 


Cephalothorax 


Abdominal segments 


Seta (3h I I Il IV 
0 — — 1 1 1 
1 2 10-14(10)* 5,6(6) 2,3(3) 2,3(2) 
2 2,3(3) 1 I 1 1 
3 23323) 1,2(1) l 1 2-4(3) 
4 2-4(3) 5—7(6) 5,6(5) 3-5(4) 2-6 
5 3 1 6-11 5—8(5) i 
6 1-3(2) 1 1 1 1 
7 2-4(3) 3 3,4(3,4) 4,6(4,6) 3-8 
8 3,4(3) = a 4-7 2 
9 1 1 1 1 1 
10 1,2(1) at 1,2(1) 120) 2 
11 3,4(3) 1 2;3(2;3) 3 1,2(2) 
12 12 @,2) = = = aa 
13 — _ _ — = 
14 -- — = == — 


* Primary branches. 
+ Presumed (see legend for Fig. 4). 
+ Alveolus only. 


specimens, 13-VI quite variable in size and 
number of branches (3—12), apparently ab- 
sent in Sa. chloropterus. Genital lobe: 
Lightly tanned, length (male only) about 
0.36 mm. Paddle: Lightly tanned, asym- 
metrical, broadest at base, tapered distally, 
inner part narrow, less than 0.5 width of 
outer part, and minutely spiculate at apex; 
length about 0.7 mm, width at widest point 
about 0.45 mm, index 1.55. 
Systematics.—Sabethes conditus resem- 
bles Sa. tridentatus and Sa. glaucodaemon, 
but differs in having distinctly shorter upper 
meseperimeral setae. These setae reach the 
lateral area of the mesopostnotum in Sa. 
conditus and extend to near the middle of 
the mesopostnotum in the other two spe- 
cies. The male of Sa. conditus also differs 
in the presence of ventral pale scaling on 
the proboscis, the pattern of pale scaling on 
the mid-tarsi, and the structure of the gen- 
italia. Sabethes conditus has undoubtedly 
been misidentified as Sa. chloropterus in 
the past, mainly because most studies have 
focused on females that appear to be essen- 
tially isomorphic for the two species. 
Knight and Stone (1977) list four junior 
synonyms for Sa. chloropterus: Sabethes 


nitidus (Theobald), Sabethoides confusus 
(Theobald), Sabethoides rangeli (Surcouf 
and Gonzales-Rincones), and Sabethoides 
imperfectus (Bonne-Webster and Bonne). 
Sabethes nitidus was included in error be- 
cause the lectotype designation by Belkin 
(1968) validated this nominal taxon as a 
distinct species of subgenus Sabethes, and 
the paralectotype male that Knight and 
Stone used as a basis for synonymy with 
Sa. chloropterus has no taxonomic status. 
We know from concurrent studies of Sa- 
bethoides that the holotype female of Sa- 
bethoides imperfectus is not conspecific 
with Sa. chloropterus and will be formally 
synonymized with another species in a fu- 
ture revision of the subgenus (by R.E.H.). 
The holotype females of Sabethoides con- 
fusus and Sabethoides rangeli are both in 
very poor condition, and even though Sa- 
bethoides confusus has no midlegs, both 
specimens clearly belong to a species of Sa- 
bethoides other than Sa. glaucodaemon or 
Sa. tridentatus. The holotype of Sabethoi- 
des rangeli has faint white scaling on the 
ventral surface of the proboscis (0.5 to 0.8 
from base), a feature that is not known in 
females of Sa. chloropterus or the pre- 


VOLUME 102, NUMBER 4 999 
Table 2. Continued. 
Abdominal segments Paddle 

Vv VI VII VII IX le 
1 1 1 1 — => 
2 2,3(2) 23 (6) —- — a 
1 1 1 — — — 
DS (2) 2,3(2) 3,4(3) a = = 
3—6(6) 2 1 1,2(1,2) = = 
1 1 3—5(4) — _ — 
1E2(2) 1252) 5,6(5,6) = wer 
5-8 1272) 1,2(1,2) — — — 
2,3(2) 6—9(7) 10—15(14) — — — 
1 1 13—15(14) 23—27(26) = = 
1 1 Asi) = = == 
1,2(2) 4-6(4) 1 — — ae 
el 12 at we ie he 


sumed females of Sa. conditus examined in 
this study. In the absence of any evidence 
to suggest that the holotype of either Sa- 
bethoides confusus or Sabethoides rangeli 
may be conspecific with Sa. conditus rather 
than Sa. chloropterus, these nominal forms 
are retained in synonymy with the latter 
species. 

The male of Sa. conditus is easily distin- 
guished from the male of Sa. chloropterus 
in having a lesser amount of pale scaling 
on the proboscis and midtarsus, and a very 
differently constructed gonostylus. The pro- 
boscis of Sa. conditus has ventral pale scal- 
ing extending from 0.5 to 0.8 beyond the 
base whereas in Sa. chloropterus it extends 
from 0.3 to 0.8 beyond the base. Similarly, 
the midtarsus has pale scaling on the ante- 
rior surfaces of tarsomeres 2—4 in Sa. con- 
ditus (also on the posterior surface of tar- 
someres 3 and 4 in the paratype from Bra- 
zil) whereas it occurs on the anterior sur- 
faces of tarsomeres 2 and 3 and the 
posterior surfaces of tarsomeres 1—4 in Sa. 
chloropterus. 

The two females of Sa. conditus from 
Brazil are presumed to belong to this spe- 
cies because specimen numbers on the la- 
bels indicate that they were collected and 
reared from pupae along with the paratype 


male. Apparently no collection records, ad- 
ditional data or associated pupal exuviae 
exist in the Smithsonian Institution to con- 
firm this (J. Pecor, personal communica- 
tion). The two females have the same pat- 
tern of pale scaling on the midlegs and ap- 
pear to resemble verified females of Sa. 
chloropterus in overall habitus (the midlegs 
of Sa. chloropterus sometimes also have a 
variable amount of pale scaling on the pos- 
terior surfaces of tarsomeres 2—4). If these 
two specimens are females of Sa. conditus, 
then the female of this species is indistin- 
guishable from the female of Sa. chlorop- 
terus. 

No diagnostic characters were found to 
distinguish the larva and pupa of Sa. con- 
ditus from those of Sa. chloropterus, but 
some partially differential characteristics 
have been noted that may prove to be useful 
once additional material becomes available 
for further study. In the pupa, seta 4-IV has 
2—6 branches whereas specimens of Sa. 
chloropterus only have 1 or 2 (usually 2), 
and seta 13-VII, a unique feature among 
mosquitoes, is present on both sides in the 
two available specimens of Sa. conditus. In 
the larva, the mean sum of branches of seta 
13-III-V on one side is 17 compared to 12 
in Sa. chloropterus, comb scales apparently 


1000 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


eee Hoey 


Fig. 4. A, B, Pupa of Sabethes (Sabethoides) conditus. A, Left side of cephalothorax, dorsal to night. B, 
Dorsal (left) and ventral (right) aspects of metathorax and abdomen. C, Localities in Venezuela and Brazil where 
type specimens of Sa. conditus were collected (see Material examined). Seta 5-[V was absent in available 
specimens and is drawn similar to 5-V,VI based on this development in Sabethoides. CT = cephalothorax; P = 
paddle; I-VIII = abdominal segments; 1—14 = setal numbers for specified areas, e.g., seta 3-I. Scale in mm. 


VOLUME 102, NUMBER 4 


lack the fringes of minute spicules that are 
present in Sa. chloropterus, and seta 4-X is 
double whereas it has 3 or 4 branches in 
Sa. chloropterus. 

Etymology.—The specific name of con- 
ditus is a Latin adjective (masculine) mean- 
ing unseen or hidden. The name refers to 
the fact that the species undoubtedly has 
been mistaken for Sa. chloropterus in the 
past. 

Bionomics.—The type specimens from 
Venezuela were collected as larvae found in 
brownish water contained in a small hole 
located 0.6 m above ground level in a le- 
guminous tree. The treehole also contained 
larvae of Culex (Anoedioporpa) conserva- 
tor Dyar and Knab. Nothing else is in def- 
initely known about the bionomics of Sa. 
conditus, but it is likely that some of the 
information published about Sa. chloropte- 
rus actually applies to this species. 

Distribution.—The type specimens were 
collected at the two localities indicated in 
Fig. 4C, one in northern Venezuela and the 
other in western Brazil. Sabethes conditus 
is undoubtedly widely distributed between 
and around these localities. Reports of Sa. 
chloropterus from this region may apply to 
Sa. conditus or a mixture of the two spe- 
cies. 

Material examined.—Eleven specimens 
(3 6,2 6 genitalia, 2 2, 2 larval exuviae, 
2 pupal exuviae), including 2 larval rear- 
ings. VENEZUELA: Aragua, Ocumare de 
la’ Costa, Puerto Ocumare, just E of 
(19PFM3459), near sea level, 10 Jul 1969, 
J. A. Bergland and J. Valencia, 1 ¢ with 
associated larval and pupal exuviae and dis- 
sected genitalia on separate slides (VZ133- 
11), associated larval and pupal exuviae of 
lost 6 (VZ133—22) and 1 6 (VZ133-1) 
(small treehole in partial shade 4 m above 
ground in leguminous tree in coconut plan- 
tation; water temporary, brown, turbid). 
BRAZIL: Rond6nia, Costa Marques, 31 Jan 
1992, USAMRU-B personnel, 1 d6 (BR513 
(65)-102) with dissected genitalia on micro- 
scope slide and 2 2 (BR513 (65)-100 and 
-102). 


1001 


The males above comprise the type series 
of Sa. conditus. The holotype (VZ133-11), 
with associated larval and pupal exuviae 
and dissected genitalia on separate micro- 
scope slides, and 3 paratypes (VZ133-1, 
VZ133-22, BR513 (65)-102) are deposited 
in the National Museum of Natural History, 
Smithsonian Institution. The two females 
are excluded from the type series because 
they are not definitely associated with the 
males, and may not be conspecific with 
them. 


ACKNOWLEDGMENTS 


We thank personnel of the Walter Reed 
Biosystematics Unit, Smithsonian Institu- 
tion, Washington, DC, for the loan of spec- 
imens. We are grateful to Robert Hancock, 
Cumberland College, Williamsburg, Ken- 
tucky, for assistance and advice to Dusty 
Moses. Jane Carter of the same college is 
acknowledged for co-ordinating the Inter- 
national Enrichment Program that provided 
the Biological Internship to Dusty Moses 
that made this study possible. 


LITERATURE CITED 


Belkin, J. N. 1968. Mosquito studies (Diptera, Culic- 
idae) IX. The type specimens of New World mos- 
quitoes in European museums. Contributions of 
the American Entomological Institute (Ann Ar- 
bor) 3(4): 1-69. 

Galindo, P., E. de Rodaniche, and H. Trapido. 1956. 
Experimental transmission of yellow fever by 
Central American species of Haemagogus and Sa- 
bethes chloropterus. American Journal of Tropical 
Medicine and Hygiene 5: 1022-1031. 

Galindo, P., E. de Rodaniche, and C. M. Johnson. 
1959. St. Louis encephalitis in Panama. 1. Isola- 
tion of the virus from forest mosquitoes and hu- 
man blood. American Journal of Tropical Medi- 
cine and Hygiene 8: 557-560. 

Hall, C. R., T. M. Howard, and R. E. Harbach. 1999. 
Sabethes (Peytonulus) luxodens, a new species of 
Sabethini (Diptera: Culicidae) from Ecuador. Me- 
morias do Instituto Oswaldo Cruz 94: 329-338. 

Harbach, R. E. 199la. A new subgenus of the genus 
Sabethes (Diptera: Culicidae). Mosquito System- 
atics 23: 1-9. 

. 1991b. Ontogeny of the larval stage of Sa- 

bethes chloropterus, with special reference to setal 

development and phylogenetic implications for 


1002 


the family Culicidae (Diptera). Mosquito System- 

atics 23: 10-24. 

. 1994. The subgenus Sabethinus of Sabethes 

(Diptera: Culicidae). Systematic Entomology 19: 

207-234. 

. 1995a. A new Sabethes of the subgenus Pey- 

tonulus (Diptera: Culicidae) with an unusual 

fourth-instar larva. Entomologica Scandinavia 26: 

87-96. 

. 1995b. Two new species of the subgenus Pey- 
tonulus of Sabethes (Diptera: Culicidae) from Co- 
lombia. Memorias do Instituto Oswaldo Cruz. 90: 
583-587. 

Harbach, R. E. and K. L. Knight. 1980. Taxonomists’ 
Glossary of Mosquito Anatomy. Plexus Publish- 
ing, Inc., Marlton, New Jersey, xi + 415 pp. 

. 1982. Corrections and additions to Taxono- 

mists’ Glossary of Mosquito Anatomy. Mosquito 

Systematics 13: 201—217. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Harbach, R. E. and J. L. Petersen. 1992. Two species 
previously confused under the concept of Sabeth- 
es tarsopus in Central America (Diptera: Culici- 
dae). Mosquito Systematics 24: 102-124. 

Harbach, R. E. and E. L. Peyton. 1991. Transfer of the 
subgenus Davismyia from Wyeomyia to Sabethes 
and description of the type species, Miamyia pe- 
trocchiae (Diptera: Culicidae). Mosquito System- 
atics 22: 149-159. 

Heinemann, S. J. and J. N. Belkin. 1978. Collection 
records of the project ““Mosquitoes of Middle 
America” 11. Venezuela (VZ); Guianas: French 
Guiana (FG, FGC), Guyana (GUY), Surinam 
(SUR). Mosquito Systematics 10: 365—459. 

Knight, K. L. and A. Stone. 1977. A Catalog of the 
Mosquitoes of the World (Diptera: Culicidae). 2nd 
edition. Thomas Say Foundation 6: ix + 1—611. 

Lane, J. 1953. Neotropical Culicidae, Vol. Hl. Univer- 
sity of SAo Paulo, Sao Paulo. pp. 1054-1098. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 1003-1009 


CORRECTIONS AND ADDITIONS TO THE “CATALOG OF 
THE STILT BUGS, OR BERYTIDAE, OF THE WORLD 
(INSECTA: HEMIPTERA: HETEROPTERA)” 


THOMAS J. HENRY AND RICHARD C. FROESCHNER 


(TJH) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. 
Department of Agriculture % National Museum of Natural History, Washington, DC 
20560-0168, U.S.A. (e-mail: thenry @sel.barc.usda.gov); (RCF) Department of Entomol- 
ogy, National Museum of Natural History, Smithsonian Institution, Washington, DC 
20560-0105, U.S.A. 


Abstract.—The catalog of the stilt bugs of the world, containing entries for 36 valid 
genera and 168 valid species, appeared in early 1998. Since then, we have discovered a 
number of errors thanks to the help of several colleagues. Corrections and additions range 
from minor misspellings to the omission of taxa. Among the more important entries 
concern the date of description of Metacanthus Costa and its type species and the addition 
of the two overlooked species, Neides propinquus Horvath and Gampsocoris gatai Giin- 


ther. 


Key Words: 
misspellings, literature cited 


The world stilt bug catalog (Henry and 
Froeschner 1998. Catalog of the stilt bugs, 
or Berytidae, of the world (Insecta: Hemip- 
tera: Heteroptera). Contributions of the 
American Entomological Institute 30(4): 1— 
72.) appeared in early 1998. Since then, we 
have discovered a number of mistakes, in- 
cluding those brought to our attention by 
Drs. I. M. Kerzhner, Jean Péricart, Wolf- 
gang Rabitsch, and J. L. Stehlik. These er- 
rors, most of which involve Old World taxa, 
have prompted us to provide this paper in 
the interest of making the corrections avail- 
able for inclusion in Dr. Péricart’s forth- 
coming catalog of the Palearctic Berytidae. 

The corrections, additions, and other 
changes in this paper are arranged by page 
and follow the general format of our Cata- 
log (Henry and Froeschner 1998). The one 
exception to the format is that we give only 
“author and date’’ for text references doc- 


Heteroptera, Berytidae, catalog, omitted taxa, gender, type fixations, dating, 


umenting the country distribution. The user 
should consult the bibliography for the full 
citation of these references. 


CORRECTIONS AND ADDITIONS 


Page 6. To the distribution of Apoplymus 
pectoralis Fieber add Albania (Josifov, 
1986) and Cyprus (Lindberg, 1948). 

Page 8. Under the genus Neides, the type 
designation for junior synonym Podi- 
cerus Duméril is not ‘Monotypic.”’ 
Correct type of designation to “‘Desig- 
nated by Kirkaldy, 1900, Entomologist, 
33(449): 265.” 

Page 8. Under entry for Neides aduncus 
Fieber, 1859, correct spelling of type lo- 
cality “‘Corisca”’ to ‘‘Corsica.”’ 

Page 8. Under distribution for Neides af- 
ghanus Seidenstiicker correct the distri- 
bution of “‘Russia’”’ to ‘Tajikistan and 
Turkmenistan.” 


1004 


Page 9. Add following species after Neides 


gomeranus Heiss: 
Neides propinquus Horvath, 1901 

1901 Neides propinquus Horvath, 
Drit. Asia. Fors:, Grafs Bug, Zich> 2: 
259. [Siberia]. 

Distribution: Mongolia, Russia (Sibe- 
rian, Far Eastern) (Hoberlandt, 1968, 
Acta Faun. Entomol. Mus. Natl. Pragae, 
13: 51; Kanyukova, 1988, In: Key In- 
sects Far-Eastern USSR, Hemiptera, 2: 
882). 

Host: Unknown. 


Page 9. Under 1865 Neides depressus entry, 


correct date given for Reuter from 1971 
to 1871. 


Page 9. Under synonymy of Neides tipu- 


larius (Linnaeus), the junior synonym 
Cimex araneoides Goeze was synony- 
mized by Harrer (G. A.), not Schaeffer. 
From distribution, transfer “‘Far East- 
ern’ records to Neides propinquus (I. 
M. Kerzhner, personal communication, 
following Kanyukova, 1988, Jn: Key In- 
sects Far-Eastern USSR, Hemiptera 2: 
882), and also drop ending “k’’ from 
‘““‘Derzhaskyk“‘and change 2™ “u’’ in 
‘“‘Lukushuk”’ to an “a.” To the distri- 
bution of N. tipularius add Greece (Jos- 
ifov 1986), Hungary (Horvath, 1898), 
Luxembourg (Reichling and Gerend, 
1994), and Slovakia (Stehlik and Vavii- 
nova 1995); and to notes of N. tipular- 
ius add ‘and Stehlik and Vavrinova 
(1995, Acta Mus. Moraviae, 79: 159) 
discussed the habitats, host, and life his- 
tory.” 


Page 11. Under host of Paleologus feanus 


and distribution of Paraberytus similis, 
correct spelling of “‘Linnavuouri’’ to 
*““Linnavuori.” 


Page 12. Authorship for the tribe Berytini- 


ni, derived from the nominate family 
name Berytinidae (based on the type ge- 
nus Berytinus Kirkaldy), should be 
credited to Southwood and Leston, 
1959, not Henry, 1997a. 


Page 12. To the distribution of Berytinus 


clavipes (Fabricius) add Bulgaria (Josi- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


fov, 1986), Hungary (Horvath, 1898), 
Luxembourg (Reichling and Gerend, 
1994), Macedonia (Géllner-Scheiding, 
1978), Romania (Sienkiewiz, 1964), 
Slovakia (Stehlik and Vavrinova, 1995), 
Slovenia (Gogala and Gogala, 1991), 
Switzerland (Rampazzi and Dethier, 
1997), and Yugoslovia (Horvath, 1903); 
and to notes of B. clavipes add ‘‘and 
Stehlik and Vavrinova (1995, Acta Mus. 
Moraviae, 79: 154) discussed the habi- 
tats, life cycle, and hosts.” 

Page 13. To the distribution of Berytinus 
consimilis (Horvath) add Slovakia 
(Stehlik and Vavirinova, 1995); and to 
notes of B. consimilis add ‘‘and Stehlik 
and Vavrinova (1995, Acta Mus. Mo- 
raviae, 79: 159) discussed the habitats 
and the host.” 

Page 13. To the distribution of Berytinus 
crassipes (Herrich-Schaeffer) add Slo- 
vakia (Stehlik and Vavfrinova, 1995, 
Acta Mus. Moraviae, 79: 159); and to 
notes of B. crassipes add ‘‘and Stehlik 
and Vavrinova (1995, Acta Mus. Mo- 
raviae, 79: 159) discussed the habitats, 
host, and wing polymorphism.” 

Page 14. To the distribution of Berytinus 
geniculatus (Horvath) add Slovakia 
(Stehlik and Vavrinova, 1995); and to 
notes of B. geniculatus add ‘and Steh- 
lik and Vavrinova (1995, Acta Mus. 
Moraviae, 79: 160) discussed the habi- 
tats, hosts, and the life cycle.” 

Page 14. To the distribution of Berytinus 
distinguendus (Ferrari) add Egypt (Lin- 
navuori, 1964) and Hungary (Horvath, 
1898). 

Page 15. Change date of publication of Ber- 
ytinus hiriticornis (Brullé) and subspe- 
cies B. hirticornis hirticornis (Brullé) 
from 1835 to 1836, following dating 
used in Palearctic Catalog (Aukema and 
Rieger, 1996); to the distribution of B. 
hirticornis add Luxembourg (Reichling 
and Gerend, 1994) and Slovakia (Steh- 
lik and Vaviinova, 1995); and add to 
notes of B. clavipes ‘‘and Stehlik and 
Vavrinova (1995, Acta Mus. Moraviae, 


VOLUME 102, NUMBER 4 


79: 157) discussed the habitats and 
hosts.” 

Page 15. To the distribution of Berytinus 
hirticornis nigrolineatus (Jakovlev) add 
Bulgaria (Josifov, 1986) and Greece 
(Rieger, 1995). 

Page 15. To the distribution of Berytinus 
hirticornis pilipes (Puton) add Greece 
(Heiss, 1984) and Spain (Wagner, 
1960). 

Page 16. To the distribution of Berytinus 
minor minor (Herrich-Schaeffer) add 
Albania and Bulgaria (Josifov, 1986), 
Austria, Romania, and Switzerland 
(Sienkiewicz, 1964), Hungary (Hor- 
vath, 1898), Luxembourg (Reichling 
and Gerend, 1994), and Slovenia (Go- 
gala and Gogala, 1991). 

Page 17. Under distribution of Berytinus 
montivagus (Meyer-Diir), drop second 
‘“*k” in Derzhansky, 1997; to the distri- 
bution of B. montivagus add Slovakia 
(Stehlik and Vavirinova, 1995); and to 
notes of B. montivagus add ‘“‘and Steh- 
lik and Vavirinova (1995, Acta Mus. 
Moraviae, 79: 161) discussed the habi- 
tats, hosts, and the life cycle.” 

Page 17. To the distribution of B. signoreti 
(Fieber) add Luxembourg (Reichling 
and Gerend, 1994), Slovakia (Stehlik 
and Vaviinova, 1995), and Tunisia (Lin- 
navuori, 1965), and to the notes of B. 
signoreti add the reference “‘Stehlik and 
Vaviinova, 1995, Acta Mus. Moraviae, 
TI 2NO2 

Page 18. Under heading of Berytinus stran- 
gulatus (Rey), correct date from 1887 
to 1888. Also, correct spelling of 
‘“LExchange”’ to ““L’Echange”’ and the 
type locality from ‘‘Europe’’ to 
wEtance, ”’ 

Page 18. To the distribution of B. striola 
(Ferrari) add Slovakia (Stehlik and Va- 
vrinova, 1995) and to notes of B. striola 
add ‘‘and Stehlik and Vavrinova (1995, 
Acta Mus. Moraviae, 79: 163) discussed 
the habitats, hosts, and life cycle.” 

Page 20. To distribution of Gampsocoris 
culicinus culicinus Seidenstiicker add 


1005 


Slovakia (Stehlik and Vavrinova, 1995); 
and to notes of G. culicinus add ‘and 
Stehlik and Vavrinova (1995, Acta Mus. 
Moraviae, 79: 165) discussed the habi- 
tats, host, and life history.” 

Page 21. To the distribution of Gampsocor- 
is enslini Seidenstiicker add Albania, 
Armenia, Azerbaidzhan, Bulgaria, 
Georgia, Greece, Iran, Yugoslavia (Per- 
icart, 1984). 

Page 21. Add following species entry: 

Gampsocoris gatai Giinther, 1996 

1996 Gampsocoris gatai Gitnther, 
Mitt. Int. Entomol. Ver., 21: 125. 
[Spain]. 

Distribution: Spain. 

Host: Teucrium charidemi Sandwith 
[Lamiaceae] (Giinther, 1998, Russian 
Entomol. Soc. 1: 106); the host was giv- 
en incorrectly as Ononis natris ssp. ra- 
mosissima (Desf.) Batt. in original de- 
scription (Giinther, 1996, Mitt. Int. En- 
tomol. Ver., 21: 128). 

Page 23. Under Gampsocoris punctipes 
punctipes (Germar) entry, correct Bur- 
meister, 1835, reference from ““Handbk” 
to “Handbch.”” Under Wagner, change 
date 1955 to 1954 and spelling of “‘sei- 
denstuckeri”’ to “‘seidenstueckeri.”’ To 
the distribution add Luxembourg 
(Reichling and Gerend, 1994), Saudi 
Arabia (Linnavuori, 1986), and Slovak- 
ia (Stehlik and Vavrinova, 1995); and to 
notes of G. punctipes add ‘‘and Stehlik 
and Vavrinova (1995, Acta Mus. Mo- 
raviae, 79: 166) discussed the habitats, 
host, and life history.” 

Page 30. Close parentheses and eliminate 
one period at end of ‘Distribution’ for 
Capyella horni Breddin. 

Page 29. Under genus Capyella and in Lit- 
erature Cited (p. 53), correct date for 
Capytum Strand from 1926 to 1928. 

Page 32. Under Jalysus albidus Stusak, cor- 
rect spelling of host genus from ‘‘Ad- 
ensotemma”’ to ‘‘Adenostemma.”’ Also 
modify spelling in host index, p. 56, un- 
der both the generic and species entries. 


1006 


Page 35. We incorrectly listed Berytus ele- 
gans Costa, 1847 (nec Curtis) as a ju- 
nior synonym of Berytus meridionalis 
Costa, 1843 and as the type species of 
Metacanthus Costa, 1847. This interpre- 
tation is in error and should have re- 
ferred to Neides elegans Curtis, as noted 
on page 23 of our catalog, where we 
correctly list Neides elegans Curtis as a 
junior synonym of Berytus punctipes 
(the type species of Gampsocoris Fuss). 
The problem leading to this situation 
began when Costa (1843) described his 
species Berytus meridionalis and sug- 
gested that it should be placed in a new 
genus, which he tentatively (but not for- 
mally) named Metacanthus. Later, he 
(Costa, 1847) described and more for- 
mally proposed the name Metacanthus, 
but used Berytus elegans Curtis (based 
on his figure) as the type species, in- 
stead of B. meridionalis. If Costa’s 
(1847) later description is followed, 
with B. elegans as the type species, then 
the genus Metacanthus becomes a ju- 
nior synonym of Gampsocoris; thus, 
disrupting the stability of the largest stilt 
bug genus. Because of this stituation, 
we fully agree with Dr. J. Péricart’s 
(personal communication) suggestion 
that Costa’s (1843) first description of 
Metacanthus should be accepted, with 
Berytus meridionalis as the type spe- 
cies, a course of action followed by him 
(Péricart, 1984) in his well-known book 
on the Berytidae of the Mediterranean 
Region. 


Therefore, modify the genus Metacan- 
thus as follows: 


Genus Metacanthus Costa, 1843 


1843 Metacanthus Costa, Cimic. Neo- 
pol., p. 26. Type species: Berytus mer- 
idionalis Costa, 1843. Original desig- 
nation. 

Note: We follow Péricart (1984), who 
considered that Costa (1843) sufficient- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ly described and validated the name Me- 
tacanthus and designated as its type, B. 
meridionalis. To accept otherwise is to 
seriously jeopardize stability and 
change the name of the largest stilt bug 
genus, Metacanthus Costa. 

Page 35. To the distribution of Metacanthus 
(Cardopostethus) annulosus (Fieber) 
add Slovakia (Stehlik and Vavrinova, 
1995): 

Page 36. Modify the subgenus Metacanthus 
as follows (following generic modifi- 
cation above, p. 35): 


Subgenus Metacanthus Costa, 1843 


1843 Metacanthus Costa, Cimic. Neo- 
pol., p. 26. Type species: Berytus mer- 
idionalis Costa, 1843. Original desig- 
nation. 

Page 36. Under Metacanthus acintus, cor- 
rect spelling of “‘Nannaizab” to ““Non- 
naizab.”’ 

Page 36. Correct spelling of species names 
Metacanthus ‘‘ataoensis” to “‘antaoen- 
sis,” three times. 

Page 36. The first combination of Gamp- 
socoris delhiensis Menon and Ghai in 
Metacanthus was by Josifov (1965, Rei- 
chenbachia 5: 285), not Wheeler and 
Schaefer (1982, Ann. Entomol. Soc. 
Am., 75: 502). 

Page 37. In note under Metacanthus linea- 
tus (Jakovlev), correct spelling of M. 
‘“‘maghrebrinus” to “‘maghrebinus.” 

Page 37. Change date for description of 
Metacanthus meridionalis (Costa) from 
1844 to 1843; delete entry for “1847 
Metacanthus elegans Costa, Cimic. 
Neapol. Cent., p. 259. [Italy]. Synony- 
mized by Bergroth, 1913, Mém. Soc. 
Entomol. Belg., 22: 177.” as explained 
under genus above (p. 35); correct spell- 
ing of “‘Corisca”’ to “‘Corsica” under 
Megalomerium pallidum entry; and cor- 
rect spelling of “‘Linnavuouri” to “Lin- 
navuori.”’ 

Page 38. In distribution under Metacanthus 
mollis Stusak, correct spelling of Lin- 
navuouri to Linnavuori; and to the dis- 


VOLUME 102, NUMBER 4 


tribution of M. mollis add Saudi Arabia, 
Somalia, and Yemen (Linnavuori, 
1986). 

Page 39. In distribution under Metacanthus 
nitidus Stusak, correct spelling of ‘‘Lin- 
navuouri”’ to ““Linnavuori.”’ 

Page 39. Change spelling of Metacanthus 
“pertenerus’’ (Breddin) to M. “‘perte- 
nerum”’ three times at species and both 
subspecies entries. Also add to index 
entries for ‘““Metacanthus pertenerum” 
(p. 68) and ‘‘Metacanthus, pertenerum 
vittatus’’ (p. 72). 

Page 40. For the type species of the genus 
Neometacanthus and under the species 
Neometacanthus picticornis, correct 
spelling of ‘‘Noualheir’” to ‘‘Noual- 
hier.” 

Page 42. To the distribution of Yemma 
gracilis Linnavuori add Yemen (Linna- 
vuori and van Harten, 1997). 

Page 43. The genus Yemma is feminine; 
therefore correct spelling of the specific 
epithet ““Yemma signatus’’ to “‘Yemma 
signata’”’ in first line and add “‘[sic]”’ to 
Hsiao, 1977 entry. 

Page 44. Under Metatropis rufescens, cor- 
rect Burmeister, 1835, reference from 
*“Handbk.”’ to “‘Handbch.” To the dis- 
tribution of M. rufescens add Luxem- 
bourg (Reichling and Gerend, 1994) 
and Slovakia (Stehlik and Vaviinova, 
1995); and to notes of M. rufescens add 
“and Stehlik and Vavrinova (1995, Acta 
Mus. Moraviae, 79: 159) discussed the 
habitats, host, and life history.” 

Page 45. The genus Metatropis is feminine; 
therefore correct spelling of the specific 
epithet “Metatropis tesongsanicus’’ to 
*““Metatropis tesongsanica’’ in first line 
and add “‘[sic]” after original combi- 
nation. 

Page 45. Correct spelling of “Reiger” to 
“Rieger” under Aukema and Rieger, 
1995 and 1996. 

Page 46. Change date of Brullé 1835 ref- 
erence to 1836, following the Aukema 
and Rieger (1996). 

Page 46. Correct dates of Costa, A. from 


1007 


1844-1862 to 1843-1862, including 
first part “‘1843 (separate, part 1, 7:143- 
405 ...,” following Kerzhner (1983, 
Mitt. Zool. Mus. Berlin 59: 191). 

Page 46. Under Curtis (1824—40) reference 
correct spelling of “‘Literatureae”’ [sec- 
ond to last line] to ‘“‘Literaturae.”’ 

Page 48. Under Germar, 1817—1847, refer- 
ence, switch order of dates from ‘‘16— 
17 (1837); 18-19 (1936)” to 1936 and 
1937; and under Gmelin, 1790, refer- 
ence, correct spelling of ‘“‘editre’’ to 
“‘edite’’; under Goeze, 1778, reference, 
capitalize “‘Natursystems.”’ 

Page 49. Under Herrich-Schaeffer refer- 
ence, change leading date 1851 to 
1836-1853 and title from ‘‘Wazenarti- 
gen Insecten” to “Die Wanzenartigen 
Insecten’’; under Horvath, 1885, delete 
“de” in Revue d’Entomologie; under 
Horvath, 1901, change ‘“‘Forsehungsre- 
ise’ to “‘Forschungsreise’’; and under 
Horvath, 1912, change “‘Tijschrift’’ to 
**Tijdschrift.”’ 

Page 50. Under Josifov, 1959, capitalize 
‘“‘art’’ in Heteropteren-Art; under Josi- 
fov, 1965, capitalize “‘systematik”’; un- 
der Josifov, 1974, change “In Russian” 
to “In Bulgarian”; under Lindberg, 
1934, change “‘F’’ to lower case in 
‘“*For’’; and under Lindberg, 1958, cor- 
rect journal spelling to Societas ““Scien- 
tiarum Fennicae.”’ 

Page 51. Under A. Puton, 1876, add to be- 
ginning of title ““Notes pour.” 

Page 52. Under Putshkov, 1974, add *‘[In 
Ukranian]’’; under Rey, 1888, change 
““L-Exchange”’ to “‘L_Echange’’; change 
authorship of J. C. Schaeffer, 1784 to 
G. A. Harrer, 1784 and move forward 
alphabetically; and under Scholtz, 1847, 
change “‘de’’ to “‘der.”’ 

Page 53. Under Stal, 1870-1876, correct 
*“Svenskaps”’ to ““Svenska’’; and under 
Strand: = 1926; cormece, the date to 
**1928,”’ change “‘nomenclatoric’’ to 
**nomenclatorica,’’ and change “‘Arkiv”’ 
to “Archiv” and insert “‘fiir.”’ 


1008 


Page 54. Under Stusak, 1976 a 1976c, spell 
out “Bulgarian Academy of Sciences.” 

Page 55. Delete Wagner 1955 “Ergebnisse, 
einer ....’’; under Wagner, 1965, cor- 
rect spelling of ‘‘Commentiones”’ to 
‘**Commentationes.”’ 


Add the following papers to the bibli- 
ography: 


Gogala, A. and M. Gogala. 1991. True bugs of Slo- 
venia (Insecta, Heteroptera). Biologiski Vestnik 
37: 11-44. 

GGllner-Scheiding, U. 1978. Beitrag zur Kenntnis 
der Heteropterenfauna Mazedoniens. Acta Musei 
Macedonici Scientiarum Naturalium 15(6): 145— 
149. 

Giinther, H. 1996. Gampsocoris gatai n. sp., eine 
neue Keulenwanzenart aus Stidspanien (Heterop- 
tera: Berytidae). Mitteilungen der Internationalen 
Entomologischen Vereins 21: 125—129. 

Giinther, H. 1998. New and rare true bugs (Heter- 
optera) from southern Spain. /n: The problems of 
Entomology in Russia. Russian Academy of Sci- 
ences, Russian Entomological Society 1: 106— 
108. 

Heiss, E. 1984. Heteropteren aus Kreta II (Insecta: 
Heteroptera). Bericht des Naturwissenschaftlich- 
Medizinischen Vereins in Innsbruck 71: 141—155. 

Hoberlandt, L. 1968. Results of 1st Mongolian-Cze- 
choslovak entomological and botanical expedition 
to Mongolia. No. 17: Heteroptera (Berytidae, Pyr- 
rhocoridae, Coreidae, Rhopalidae, Acanthosoma- 
tidae, Cydnidae, Scutelleridae and Pentatomidae). 
Acta Faunistica Entomologica Musei Nationalis 
Pragae 13: 51—60. 

Horvath, G. 1898. Ordo Hemiptera. In: Fauna Regni 
Hungarieae: 1—64. Budapest. 

Horvath, G. 1903. Szerbia Hemiptera Faunaja. (Fau- 
na Hemipterorum Serbiae). Annales Musei Na- 
tionalis Hungarici 1: 1-8. 

Josifov, M. 1986. Verzeichnis der von der Balkan- 
halbinsel bekannten Heteropterenarten (Insecta, 
Heteroptera). Faunistische Abhandlungen Staa- 
tliches Museum fiir Tierkunde Dresden 14: 61—93. 

Kerzhner, I. M. 1983. Nachtrag zu den Publikations- 
daten der Separatausgabe de Werkes “‘Cimicum 
Regni Neapolitani. Centuriae’’ von A. Costa. Mit- 
teilungen aus dem Zoologischen Museum in Ber- 
lin 597 L911" 

Kirkaldy, G. W. 1900. On the nomenclature of the 
genera of the Rhynchota, Heteroptera and auchen- 
orrhynchous Homoptera. The Entomologist 
33(449): 262-265. 

Lindberg, H. 1948. On the insect fauna of Cyprus. 
Results of the expedition of 1939 by Harald Hak- 
an and P. H. Lindberg, I-II. Commentationes Biol- 
ogicae 8 (8): 1-32. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Linnavuori, R. 1964. Hemiptera of Egypt, with re- 
marks on some species of the adjacent Eremian 
region. Annales Zoologici Fennici 1: 306—356. 

Linnavuori, R. 1965. Studies on the South-and East- 
mediterranean hemipterous fauna. Acta Entomo- 
logica Fennica 21: 1—69. 

Linnavuori, R. 1986. Heteroptera of Saudi Arabia. 
Fauna of Saudi Arabia. 8: 31—197. 

Linnavuori, R. and A. van Harten. 1997. Note on 
Heteroptera (Insecta: Hemiptera) of Yemen. Fauna 
of Saudi Arabia. 16: 169-236. 

Rampazzi, E and M. Dethier. 1997. Gli eterotteri 
(Insecta: Heteroptera) delle torbiere a sfagni del 
Cantone Ticino e de Moesano (Val Calanca e Val 
Mesolcina—GR), Svizzera. Mitteilungen der 
Schweizerischen Entomologischen Gesellschaft 
70: 419-439. 

Reichling, L. and R. Gerend. 1994. Liste des Hét- 
éropteres du Grand-Duché de Luxembourg. Bul- 
letin de la Société des Naturalistes Luxembour- 
geois 95: 273-286. 

Rieger, C. 1995. Die Fauna der Agiis-Insel Santorin. 
Teil 9. Heteroptera. Stuttgarter Beitraege zur Na- 
turkunde, Ser. A (Biologie) 520: 1—26. 

Sienkiewicz, I. 1964. The catalogue of the “A. L. 
Montandon collection” of Palaearctic Heteroptera 
preserved in the “Grigore Antipa’”’ Museum of 
Natural History, Bucharest. Bucharest, 146 pp. 

Stehlik, J. L. and I. Vaviinova. 1995. Results of the 
investigations on Heteroptera in Slovakia made by 
the Moravian Museum. Acta Musei Moraviae, 
Sci. Nat. 79: 149-168 (1994). 

Wagner, E. 1954. Gampsocoris seidenstuekeri nov. 
spec. Nachrichten des Naturwissenschaftliche 
Museums der Stadt Aschaffenburg 45: 37—40. 

Wagner, E. 1960. Beitrag zur Heteropteren-Fauna 
Nordost-Spaniens. Trabajos del Museo de Zoolo- 
gia, Nueva Serie Zoologica 2(3): 3—26. 


ACKNOWLEDGMENTS 


We are grateful to I. M. Kerzhner (Zoo- 
logical Institute, Russian Academy of Sci- 
ences, St. Petersburg) for his careful review 
of the catalog and for noting the inadvertent 
omission of Neides propinquus; J. Péricart 
(Montereau, France) for his comments on 
the serious type-species problem involving 
the genus Metacanthus; J. L. Stehlik (Dept. 
Entomology, Moravian Museum, Preslova, 
Czech Republic) for pointing out an impor- 
tant paper on the stilt bugs of Slovakia; and 
to W. Rabitsch (Dept. of Ecophysiology, In- 
stitute of Zoology, Vienna, Austria) for not- 


VOLUME 102, NUMBER 4 1009 


ing numerous omitted country records and Vandenberg, Systematic Entomology Lab- 
for bringing to our attention the description oratory, U. S. Department of Agriculture, 
of the recently described Gampsocoris ga- Washington, DC, for reviewing the manu- 
tai. We also thank D. R. Smith and N. J. script and offering other useful suggestions. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 1010-1013 


THREE NEW SPECIES OF DIPOGON FOX (SUBGENUS DIPOGON) 
(HYMENOPTERA: POMPILIDAE) FROM CENTRAL AND WESTERN 
NORTH AMERICA 


HOWARD E. EVANS 


Department of Bioagricultural Science and Pest Management, Colorado State Univer- 


sity, Fort Collins, CO 80523-1177, U.S.A. 


Abstract.—Three new species are described in the genus Dipogon, subgenus Dipogon 
(Pompilidae). They are: D. kiowa (Prowers and Delta counties, Colorado), D. kenza 
(Douglas Co., Kansas), and D. anasazi (Montezuma Co., Colorado). 


Key Words: 


Species of Dipogon Fox are not often 
taken by conventional collecting methods, 
as they are rarely attracted to flowers or 
honeydew. They occur principally in wood- 
ed areas, where most nest in cavities in 
wood. Males are rarely taken except when 
reared from trap nests (Medler and Koerber 
1957, Krombein 1967). One species of sub- 
genus Deuteragenia Sustera has been 
reared from stems of weeds and garden 
plants (Williams 1966) and a species of 
subgenus Dipogon has been reared from 
stems of Sambucus (Wasbauer 1966), but 
most specimens have been taken in asso- 
ciation with trees. Townes (1957) recog- 
nized five species of subgenus Dipogon 
north of Mexico, but four more have since 
been added (Wasbauer 1960, 1966; Evans 
1987). Three more species are added here, 
all thus far known only from females. All 
have the frons and thoracic dorsum pol- 
ished and largely devoid of recumbent pu- 
bescence and the punctures of the frons dis- 
tinct but minute and widely spaced. Thus 
they belong to the graenicheri group of 
Townes (1957) provided the limits of that 
group are expanded to include species in 
which the microtrichiae of the fore wing are 
not necessarily much larger and more 


Hymenoptera, Pompilidae, Dipogon, new species, North America 


crowded in the fuscous bands than else- 
where. 


Dipogon (Dipogon) kiowa Evans, 
new species 


Holotype.—?, COLORADO: Prowers 
Co., Lamar, tree trunks, 4—7 Sept. 1998 (D. 
Leatherman) [National Museum of Natural 
History]. 

Paratypes.—, same data; 2, same lo- 
cality and collector, Willow Creek Park, 
15-16 Aug. 1998; 6 2, same locality and 
collector as type, 26-31 July 1999; 9, 
COLORADO: Delta Co., Crawford State 
Park, ex Utah juniper, 5 Aug. 1999 (D. 
Leatherman) [Paratypes in National Muse- 
um of Natural History and Colorado State 
University]. 

Etymology.—Named for the Kiowa In- 
dians that formerly roamed the Arkansas 
Valley. On July 27, 1820, members of the 
Long Expedition visited an encampment of 
Kiowas not far from the type locality. 

Description of type.—Length 5.5 mm, 
fore wing length 4.2 mm. Head ferruginous, 
including mouthparts and antenna, except 
first two and last antennal segments par- 
tially infuscated, ocellar triangle and paired 
streaks just below them weakly infuscated. 


VOLUME 102, NUMBER 4 


Pronotum ferruginous, with a pair of fus- 
cous spots anterodorsally; remainder of tho- 
racic dorsum and central part of propodeum 
black; meso- and metapleura and lateral 
parts of propodeum ferruginous; thoracic 
venter black. Metasoma black except sides 
of first tergite stained with reddish. Legs 
ferruginous except suffused with black as 
follows: upper surface of mid and hind cox- 
ae and trochanters, basal and apical extrem- 
ities of fore and mid femora and much of 
mesal surface of hind femur, outer surface 
of fore and mid tibiae and all of hind tibia, 
basal segment of mid and hind tarsi. Wings 
hyaline, with a narrow brown band at the 
basal vein and a large brown area extending 
from the stigma through the third discoidal 
cell; microtrichiae slightly denser in the 
dark bands than elsewhere, especially dense 
at the basal vein. 

Surface of head polished, only very 
faintly alutaceous; clypeus with several 
strong setae; frons with small punctures 
separated by 5—10 times their own diame- 
ters, each giving rise to a very short, erect 
seta. Thorax with integument shining, me- 
soscutum slightly more alutaceous and 
more close punctate than frons; scutellum 
and metanotum with dense, recumbent sil- 
very to golden pubescence; meso- and me- 
tapleura polished, very weakly alutaceous; 
mid and hind coxae strongly silvery-seri- 
ceous; propodeum polished, with sparse, 
small punctures; metasoma sparsely sil- 
very-sericeous. 

Width of head 1.05 times length of head 
to margin of clypeus; middle interocular 
distance 0.60 times width of head; eyes 
convergent above, upper interocular dis- 
tance 0.75 times lower. Postocellar and 
ocello-ocular distances subequal. First four 
antennal segments in a ratio of 17:9:29:22. 
Posterior margin of pronotum arcuate; slope 
of propodeum low and even, midline not 
impressed. Second submarginal cell maxi- 
mum length 3 times its maximum height 
and 1.6 times maximum length of third sub- 
marginal cell. 

Variation—Two of the paratypes are 


1011 


considerably smaller than the type, with 
fore wing length 3.2 mm. Otherwise, there 
is little variation in size or morphology 
within the series. Two of the Lamar para- 
types have the frons wholly ferruginous, 
and one Lamar paratype as well as the one 
from Delta Co. have the frons as well as 
the vertex and occiput black. These darker 
specimens also have the thorax and legs 
more heavily infuscated than in the other 
paratypes. 

Remarks.—The polished and nearly bare 
integument, combined with the contrasting- 
ly dense pubescence on the scutellum and 
metanotum, plus the unusual color pattern, 
distinguish this species. In contrast to 
graenicheri Banks (1939), the body is more 
extensively marked with black and the me- 
tasoma wholly black; also the fasciae of the 
fore wing are less intense and the microtri- 
chiae only slightly more dense within the 
wing bands. Dipogon parkeri Wasbauer 
(1966), described from Nevada, has a color 
pattern not unlike kiowa, but the head, tho- 
racic dorsum, and propodeum are duller, 
with stronger surface sculpturing and an 
overlay of appressed pubescence. In parkeri 
the eyes are less convergent above, the up- 
per interocular distance measuring 0.89 
times the lower. 


Dipogon (Dipogon) konza Evans, 
new species 


Holotype-—?, KANSAS: Douglas Co., 
Lawrence, 25 Aug. 1986, Douglas Yanega, 
on Gonolobus vine [Snow Entomological 
Museum, University of Kansas]. 

Paratypes.—3 2, same data except 26 
Aug., 4 and 27 Sept. 1896; 2, Lakeview, 
near Lawrence, 24 Sept. 1966, G. C. Eick- 
wort; 2, Breidenthal Reserve, 2 mi. N 
Baldwin, Douglas Co., 22 Aug.—8 Sept. 
1982, malaise trap, D. B. Wahl [paratypes 
at University of Kansas, Colorado State 
University, and National Museum of Nat- 
ural History]. 

Etymology.—Named for the Konza In- 
dians, using the spelling employed by early 


1012 


explorers in what is now the state of Kan- 
sas. 

Description of type.—Length 4.2 mm; 
fore wing length 3.4 mm. Body entirely 
glossy black except clypeus and sides of 
pronotum brownish; mandible dark brown 
basally, yellow-brown apically; antenna 
wholly light ferruginous, legs also of this 
color except coxae suffused with brown. 
Wings hyaline, somewhat luteous, veins on 
outer part translucent; fore wing unbanded 
but with a small brown cloud just distad of 
stigma; microtrichiae of uniform size and 
distribution except somewhat more crowd- 
ed along basal vein. 

Clypeus with several strong setae. Frons 
and vertex strongly polished, without sur- 
face sculpturing, with minute punctures 
separated by many times their own diame- 
ters, each giving rise to a short, erect seta. 
Thorax and propodeum similarly polished 
and sparsely punctate except scutellum fine- 
ly, closely punctate and mesosternum with 
fine, appressed, silvery pubescence. Meta- 
soma polished, very weakly alutaceous, 
with the usual strong setae ventrally and 
apically; first 3 tergites with only scattered, 
short setae. 

Width of head 1.06 times length of head; 
middle interocular distance 0.61 times head 
width; eyes weakly convergent above, up- 
per interocular distance 0.9 times lower. 
Vertex weakly elevated above eye tops, 
postocellar and ocello-ocular distances sub- 
equal. First 4 antennal segments in a ratio 
of 17:9:20:17. Posterior margin of prono- 
tum arcuate; slope of propodeum low and 
even, midline not impressed. Second sub- 
marginal cell with its maximum length 3 
times the maximum height, 1.5 times max- 
imum length of third submarginal. 

Variation.—Individuals in this series 
vary in fore wing length from 3.0 to 3.8 
mm. All have the deep brown to black 
glossy integument and the light ferruginous 
antennae and legs (at least beyond the cox- 
ae), but the largest female is streaked with 
ferruginous along the sides of the thorax 
and propodeum. Although there are no dis- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


tinct wing fasciae in any of the specimens, 
in two there is weak clouding along the bas- 
al vein. 

Remarks.—The clear wings, with only 
faint localized clouding in some specimens, 
set this species apart from other species of 
Dipogon except for fulleri Krombein 
(1962), a member of the subgenus Winne- 
manella. The glossy black integument con- 
trasting to the pale appendages also char- 
acterizes this species. 


Dipogon (Dipogon) anasazi Evans, 
new species 


Holotype.—?, COLORADO: Montezu- 
ma Co., Soda Canyon, Mesa Verde Nation- 
al Park, 6 Aug. 1999, B. Kondratieff, S. 
Wells, W. Cranshaw, P. Pineda, & W. Paint- 
er [National Museum of Natural History]. 

Etymology.—This species is named for 
the Anasazi, the remains of whose homes 
provide the major treasures of Mesa Verde 
National Park. 

Description of type.—Length 4.8 mm; 
fore wing length 4.4 mm. Entire body and 
appendages light ferruginous, eyes and 
ocelli contrastingly nearly black. Wings hy- 
aline, slightly luteous basally; fore wing 
with a weak brown band at basal vein and 
a broad brown cloud below the stigma and 
basal half of marginal cell. Microtrichiae of 
fore wing distinctly darker and more 
crowded within the two dark bands than 
elsewhere. 

Integument of head, thorax, and propo- 
deum strongly polished and without surface 
sculpturing, also without appressed pubes- 
cence. Clypeus with several strong setae, 
but frons with only short, erect hairs arising 
from small punctures separated by many 
times their own diameters. Thorax and pro- 
podeum similarly with sparse, small punc- 
tures except posterior fifth of mesoscutum 
and all of scutellum and metanotum with 
dense, fine punctures. Metasoma also pol- 
ished but with rather dense, fine punctures, 
posterior segments with usual stiff bristles. 

Head 1.04 times as wide as high, vertex 
roundly elevated above eye tops. Middle in- 


VOLUME 102, NUMBER 4 


terocular distance 0.62 times head width; 
upper interocular distance 0.84 times lower. 
Ocelli in a compact triangle, ocello-ocular 
distance 1.2 times postocellar distance. First 
4 antennal segments in a ratio of 19:10:25: 
21. Second submarginal cell with its max- 
imum width 2.8 times maximum height, 1.3 
times maximum width of third submarginal 
cell. 

Remarks.—This species has many fea- 
tures in common with graenicheri Banks 
(1939). It differs in the wholly pale color- 
ation and weaker wing bands; also the body 
is even more highly polished, the vertex 
more elevated above the eye tops, and the 
punctures of the frons and thorax are more 
sparse. 

The type and only known specimen was 
collected by Samuel Wells by beating ju- 
nipers in a relatively dry canyon that had 
been burned over a few years earlier. 


DISCUSSION 


Dipogon diablo Wasbauer (1960) was as- 
signed by the describer to the graenicheri 
group, but the head and thoracic dorsum 
and dull and minutely granulo-reticulate as 
well as sparsely covered with appressed pu- 
bescence. Thus it does not appear closely 
related to the species considered here. The 
following couplets may serve to separate 
females of the graenicheri group as defined 
here. 


KEY 


1. Fore wing not fasciate (may have a small 
brown spot distad of stigma); body glossy 
black, legs and antenna light ferruginous (body 
may be streaked with ferruginous along sides 
@laineSOSOMaA) wreak sehek el eek okene konza, 0.sp. 

— Fore wing bifasciate; color not as above .... 2 

2. Entire body pale ferruginous, with contrasting 
dark eyes and ocelli; integument highly pol- 
ished; vertex roundly elevated above eye tops 
RSE ese. dks ao e4 anasazi, N. sp. 

— Not entirely ferruginous nor integument as 
highly polished; vertex more weakly elevated 
ADOWUS CVS LO) SSeS Ske now closes oe oeaeeee 3 

3. Mesosoma and at least basal third of metasoma 
ferruginous; legs ferruginous 


g. graenicheri Banks 


1013 


— Mesosoma fuscous at least dorsally, metasoma 
wholly black; legs in large part fuscous .... 4 
4. Body black, antenna, fore leg, and tarsi except 
basally, tinged with fulvous; microtrichiae of 
fore wing much darker and more crowded in 
fasciae than elsewhere; scutellum and metan- 
otum not more densely pubescent than meson- 
(ORDtCH Ss wah oo Gd edict graenicheri atratus Townes 
— Mesosoma black, pleura and lateral parts of 
propodeum ferruginous; microtrichiae of fore 
wing barely darker and more crowded in fas- 
ciae than elsewhere; scutellum and metanotum 
Genselyapubescentn sas aaa Kiowa, nN. sp. 
ACKNOWLEDGMENTS 


For the loan of types and other speci- 
mens, I am indebted to Robert W. Brooks, 
University of Kansas; Michael S. Kelley, 
Museum of Comparative Zoology; Steve 
Heydon, University of California at Davis; 
and Wojciech Pulawski and Bob Zuparko 
of the California Academy of Sciences. 


LITERATURE CITED 


Banks, N. 1939. Notes and descriptions of native 
Psammocharidae. Canadian Entomologist 71: 
225-231. 

Evans, H. E. 1987. The genus Dipogon (Hymenoptera: 
Pompilidae) in the Rocky Mountains. Entomolog- 
ical News 98: 41—45. 

Krombein, K. V. 1962. Natural history of Plummers 
Island, Maryland. XIII. Descriptions of new wasps 
from Plummers Island, Maryland, (Hymenoptera: 
Aculeata). Proceedings of the Biological Society 
of Washington 75: 1-18. 

. 1967. Trap-nesting wasps and bees: Life his- 
tories, nests, and associates. Smithsonian Press, 
Washington, DC, 570 pp. 

Medler, J. T. and T. W. Koerber. 1957. Biology of Di- 
pogon sayi Banks (Hymenoptera, Pompilidae) in 
trap-nests in Wisconsin. Annals of the Entomo- 
logical Society of America 50: 621—625. 

Townes, H. 1957. Nearctic Wasps of the Subfamilies 
Pepsinae and Ceropalinae. Bulletin of the United 
States National Museum 209. 286 pp. 

Wasbauer, M. S. 1960. Taxonomic and distributional 
notes on some western spider wasps (Hymenop- 
tera: Pompilidae). Pan-Pacific Entomologist 36: 
171-177. 

. 1966. A new spider hunting wasp of the sub- 
genus Dipogon from western Nevada (Hymenop- 
tera: Pompilidae). Proceedings of the Biological 
Society of Washington 79: 17—20. 

Williams, E X. 1966. A spider-hunting wasp found 
nesting in hollow plant stems (Hymenoptera: 
Pompilidae, Pepsinae). Wasmann Journal of Bi- 
ology 24: 33-47. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 1014-1069 


CATALOGUE OF THE TYPE SPECIMENS OF TORTRICIDAE 
(LEPIDOPTERA) IN THE COLLECTION OF THE NATIONAL MUSEUM OF 
NATURAL HISTORY, SMITHSONIAN INSTITUTION, WASHINGTON, D.C. 


JOHN W. BROWN AND JON LEwIs 


Systematic Entomology Laboratory, USDA, PSI, Agricultural Research Service, % 
National Museum of Natural History, Washington, DC 20560-0168, U.S.A. (e-mail: 
jbrown @sel.barc.usda.gov) 


Abstract.—Type specimens of the family Tortricidae deposited in the collection of the 
National Museum of Natural History, Smithsonian Institution, are listed alphabetically by 
species (or subspecies), along with an abbreviated literature citation to the original de- 
scription and collection data for the specimen. The type collection includes 1,001 name- 
bearing types of Tortricidae: 851 holotypes, 50 lectotypes (6 of which are designated 
herein), 60 species represented by one or more syntypes, 38 species represented by the 
only known extant specimen from the original series, 1 neotype, and 1 “‘pseudo-type.” 
The collection includes the vast majority of the tortricid species described by C. Fernald, 
A. Busck, C. Heinrich, J. Clarke, A. Blanchard, and A. Kawabe. It also includes the types 
of numerous species described by Lord Walsingham, W. Kearfott, H. Dyar, N. Obraztsov, 


J. Razowski, J. Powell, W. Miller, R. Brown, and J. Brown. 


Key Words: 


The type collection of Tortricidae in the 
National Museum of Natural History 
(USNM), Smithsonian Institution, repre- 
sents a significant worldwide resource for 
researchers interested in tortricid moths 
(i.e., the leafrollers). The strengths of the 
collection lie in the faunas of North and 
South America, the Orient (Japan, Taiwan, 
Philippine Islands), and Oceania. The C. H. 
Fernald Collection, purchased in 1924 
(Clarke 1974), formed the foundation, con- 
taining types (primarily “‘cotypes’’) of nu- 
merous species described by Fernald, Lord 
Walsingham, and W. D. Kearfott, and a few 
described by A. Grote, B. Clemens, and A. 
Packard. Walsingham sent Fernald “‘coty- 
pes” of most of the species he described 
from California and Oregon (Walsingham 
1879) from material in The Natural History 
Museum (formerly British Museum of Nat- 


leafrollers, lectotype, holotype, syntype, original description, host plants 


ural History). Kearfott also gave Fernald 
examples from his type series; the majority 
of Kearfott’s material eventually was de- 
posited in the American Museum of Natural 
History, New York. Klots (1942) resolved 
most of the nomenclatorial difficulties as- 
sociated with the Kearfott “‘cotypes,”’ des- 
ignating numerous lectotypes, primarily in 
the collection of the American Museum of 
Natural History. Miller (1970) resolved 
most of nomenclatorial problems associated 
with the Fernald ‘‘cotypes,” designating 
lectotypes for the vast majority of his spe- 
cies of North American Olethreutinae. Few 
lectotypes have been designated for the 
Walsingham (1879) species from California 
and Oregon. 

From 1905-1930 August Busck and Carl 
Heinrich, both U.S. Department of Agri- 
culture employees at the museum, added 


VOLUME 102, NUMBER 4 


numerous tortricid types, primarily from 
North and South America, through their 
prolific taxonomic research on microlepi- 
doptera. In particular, over 150 species were 
described by Heinrich (1923, 1926) in his 
revisions of the Olethreutinae of North 
America. Also during the early part of the 
century, Walsingham’s (1914) contribution 
to the “‘Biologia Centrali-Americana”’ ap- 
peared; it included the descriptions of many 
new species, with most of the material split 
between The Natural History Museum and 
the USNM. By this time, Walsingham was 
designating a type for each new species, so 
there is little ambiguity regarding these pri- 
mary types. 

In the 1960s, revisions of New World 
genera by Obraztsov (1961, 1962, 1963, 
1964, 1966a, b, c) resulted in the addition 
of many tortricid types to the USNM col- 
lection. From 1968—1982 specific large re- 
search projects such as Alexis Diakonoff’s 
studies of the microlepidoptera of the Phil- 
ippine Islands (Diakonoff 1968) and of Sri 
Lanka (Diakonoff 1982), and J. E G. 
Clarke’s work on Neotropical ‘‘Phaloni- 
idae”’ (Clarke 1968) and the microlepidop- 
tera of Rapa Island and Micronesia (Clarke 
1971, 1976) added many types of Tortrici- 
dae. The Blanchard Collection, donated in 
1985 (Davis 1985), also added many tortri- 
cid types, nearly all from Texas. The At- 
sushi Kawabe collection, composed almost 
exclusively of Oriental species and donated 
in 1990 (Davis 1996), added significant 
numbers to the tortricid type collection as 
well, broadening greatly the geographic 
coverage. Use of the tortricid holdings of 
the USNM by specialists worldwide (e.g., 
Richard L. Brown, John W. Brown, John B. 
Heppner, William E. Miller, Jerry A. Pow- 
ell, Josef Razowski, and others) has contin- 
ued to increase type holdings. 

Through recent curatorial efforts, the 
type specimens of the family Tortricidae 
have been organized into a single collec- 
tion. The specimens are arranged in alpha- 
betic sequence by species (or subspecies) in 
individual, labeled unit trays and stored in 


1015 


13 insect drawers in the USNM. The pur- 
pose of this paper is to present a list of the 
type specimens. The list is arranged alpha- 
betically by species (or subspecies), fol- 
lowed by the author, year of publication, 
and genus (or genus and species for sub- 
species) in which the taxon was originally 
described. This information is followed by 
an abbreviated citation of the publication in 
which the description appears. The sex, 
type designation (i.e., holotype, lectotype, 
syntype, “‘type’’, neotype, ““‘pseudo-type’’), 
and collection data also are included. We 
attempt to provide the maximum collection 
data, relying on both the text of the original 
description and information on specimen la- 
bels. For example, country, county, prov- 
ince, etc. frequently are not present on the 
specimen labels; this information is provid- 
ed where retrievable accurately. Abbrevia- 
tions used in the list include the following: 
AMNH = American Museum of Natural 
History, New York; BMNH = The Natural 
History Museum, London; NHMW = Na- 
turhistorisches Museum, Wien; r.f. = reared 
from; em: = emerged; mi = mile(s); ' = 
feet (elevation). Where the collecting date 
can have more than one interpretation, it is 
presented in quotes. For example, “‘Aug. 8/ 
15”’ could be interpreted as 8 August 1915 
or 8—15 August [no year]. 

The modern practice of designating a ho- 
lotype was not followed by most tortricid 
workers before 1900 (e.g., Riley, Grote, 
Zeller), and some early authors (e.g., Wal- 
singham, Dyar) labeled all specimens of the 
original series as “‘type.’’ Where the origi- 
nal type series consists of a single speci- 
men, as clearly stated by the author, we re- 
fer to it as the holotype even though not 
specifically designated as such in the orig- 
inal description. Where only a single spec- 
imen is presumed still to be extant from the 
original series (and a type was not specifi- 
cally designated), we refer to this specimen 
as the “‘type’’ (in quotes). In this case, the 
specimen is a potential lectotype because it 
may be the only remaining representative 
from the original series. Where two or more 


1016 


specimens of the original series are present 
and no type was specifically designated in 
the original description, we refer to these 
specimens as syntypes. We defer the latter 
two cases to the future work of specialists 
to designate appropriate lectotypes for 
these, as needed. Six specimens are unpub- 
lished lectotypes unambiguously selected 
and labeled by Obraztsov (n = 3), Powell 
(n = 2), and Clarke (n = 1); these selec- 
tions are designated in this paper. 

The USNM tortricid type collection in- 
cludes 851 holotypes, 50 lectotypes (6 of 
which are designated herein), 60 species 
represented by one or more syntypes, 38 
“types” (1.e., the only known extant spec- 
imen from the original series), 1 neotype, 
and 1 “‘pseudo-type.”’ 


LIST OF SPECIES 


abiephaga Issiki, 1962 (Ariola); in Issiki 
and Mutuura, Publ. Entomol. Lab. Univ. 
Osaka Pref. 7: 3. Holotype ¢, Japan, 
Honsyu, Koozuke, Manza, 27 July 
1958, S. Issiki & T. Yasuda. 

abietana Fernald, 1908 (Argyroploce); 
Canad. Entomol. 40: 349. Holotype <6, 
USA, Massachusetts, Hampshire Co., 
Amherst. According to Fernald this spe- 
cies was “Described from three male 
specimens taken in Amherst, Mass., one 
of which I make the type .. .”’ 

abievora Issiki, 1961 (Epagoge); in Issiki 
and Mutuura, Microlepidoptera injuri- 
ous to coniferous plants in Japan: 34. 
Holotype 6, Japan, Osaka, Ikeda, 20 
November 1958, T. Kodama. 

abornana Busck, 1939 (Lorita); Bull. 
South. Calif. Acad. Sci. 38: 101. Holo- 
type 2, USA, California, Los Angeles 
Co., El Segundo, rf. Cuscuta californi- 
ca, em: 11 July 1938, W. Pierce. 

abbreviatana Walsingham, 1879 (Paedis- 
ca); Ill. Lepid. Heter. British Museum 
4: 54. Syntype ¢, USA, Pennsylvania. 
According to the original description, 
this species was described from two 
males from Washington [D.C. according 
to Heinrich 1923], August 1872. The 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


other specimen is presumed to be in the 
BMNH. The USNM specimen bears the 
typical Walsingham “Type”’ label, but 
the collecting locality is not consistent 
with the original description. 

abundantia Clarke, 1976 (Heleanna phys- 
alodes); Insects of Micronesia 9: 15. 
Holotype 3, Micronesia, Kusaie, Pu- 
kusrik, | m, 3 April 1953, J. Clarke. 

accessa Heinrich, 1931 (Epinotia); Proc. 
U.S. Natl. Mus. 79: 9. Holotype 6, Pan- 
ama, Trinidad River, March 1912, A. 
Busck. 

achlyoptera Clarke, 1976 (Cryptaspasma); 
Insects of Micronesia 9: 58. Holotype 
?, Micronesia, Guam, Ritidian, 7 Au- 
gust 1945, J. Gressitt. 

acrocroca Diakonoff, 1982 (Acanthoclita); 
Zool. Verhandel. (Leiden) 193: 28. Ho- 
lotype 6, Sri Lanka, Ratnapura District, 
Uggalkaltota, 350’, Irrigation Bunga- 
low, 31 January—8 February 1970, D. 
Davis & B. Rowe. 

acrospodia Diakonoff, 1982 (Spanistoneu- 
ra); Zool. Verhandel. (Leiden) 193: 9. 
Holotype 6, Sri Lanka, Kandy District, 
1,800’, Peak View Motel, 15—24 Janu- 
ary 1970, D. Davis & B. Rowe. 

adana Heinrich, 1923 (Rhyacionia); Bull. 
U.S. Natl. Mus. 123: 18. Holotype 6, 
USA, Massachusetts, Essex Co., Forest 
Hill, 5 April 1910, W. Raff. 

ademonia Clarke, 1968 (razona); Proc. 
U.S. Natl. Mus. 125: 42. Holotype ?, 
Costa Rica, La Florida, 500’, W. 
Schaus. 

adjuncta Heinrich, 1924 (Gypsonoma), J. 
Wash. Acad. Sci. 14: 389. Holotype 6, 
Canada, Ontario, Toronto, June 1913, 
Parish. 

adobe Brown, 1992 (Catastega); J. New 
York Entomol. Soc. 100: 224. Holotype 
6, USA, Arizona, Cochise Co., Chiri- 
cahua Mountains, 4 mi W Portal, 
5,300’, 3-6 August 1964, D. Davis. 

adoceta Diakonoff, 1964 (Bactra); Zool. 
Verhandel. (Leiden) 70: 11. Holotype 
6, Brazil, Parana, Castro, W. Schaus. 

aechnemorpha Diakonoff, 1982 (Gypsono- 


~VOLUME 102, NUMBER 4 


ma); Zool. Verhandel. (Leiden) 193: 48. 
Holotype 6, Sri Lanka, Ratnapura Dis- 
trict, Uggalkaltota, 350’, Irrigation Bun- 
galow, 31 January—8 February 1970, D. 
Davis & B. Rowe. 

aelina Diakonoff, 1982 (Cydia); Zool. Ver- 
handel. (Leiden) 193: 21. Holotype <4, 
Sri Lanka, Ratnapura District, Uggal- 
kaltota, 350’, Irrigation Bungalow, 31 
January—8 February 1970, D. Davis & 
B. Rowe. 

aenigmana Powell, 1964 (Acleris); Univ. 
Calif. Publ. Entomol. 32: 99. Holotype 
36, USA, California, Nevada Co., Truck- 
ce Och 8-15.” 

aequilibra Diakonoff, 1982 (Aemulatrix); 
Zool. Verhandel. (Leiden) 193: 3. Ho- 
lotype 6, Sri Lanka, Ratnapura District, 
Uggalkaltota, 350’, Irrigation Bunga- 
low, 31 January—8 February 1970, D. 
Davis & B. Rowe. 

aesculana Riley, 1881 (Proteopteryx); 
Trans. St. Louis Acad. Sci. 4: 321. 
“Type”? 6, USA, Missouri, “boring the 
tender terminal twigs of Buckeye and of 
Maple ...’’ Riley described this species 
from eight specimens; we have been 
able to locate only one. The specimen 
is clearly labeled “‘Type”’ in Riley’s 
hand. 

aethalea Obraztsov, 1964 (Proeulia); Proc. 
U.S. Natl. Mus. 116: 188. Holotype ¢, 
Chile, Santiago, La Obra, October 1952, 
Pena. 

aglaia Clarke, 1955 (Orthocomotis); Trans. 
Royal Entomol. Soc. London 107: 149. 
Holotype 2, Brazil, Rio de Janeiro, Ita- 
tiaya, 700 m, 12 October 1926, J. Zikan. 

agricolana Walsingham, 1879 (Paedisca); 
Ill. Lepid. Heter. British Museum 4: 42. 
Two syntypes (2 66), USA, California. 
This species was described from five 
males from California and Oregon, col- 
lected in May and June. As is typical of 
Walsingham specimens from this era, 
both of the syntypes cited above are la- 
beled ‘“‘Type.’’ Neither has an abdomen; 
although there is a slide label attached 


1017 


to one of the specimens, the slide has 
not been found. 

ainslieana Obraztsov, 1962 (Anopina); Am. 
Mus. Novit. 2082: 34. Holotype ¢, 
USA, New Mexico, Dona Ana Co., 
Mesilla, C. Ainslie. 

aktita Miller, 1978 (Rhyacionia); in Powell 
and Miller, U.S. Dept. Agric., Agric. 
Handb. 514: 24. Holotype ¢, USA, 
New Jersey, Ocean Co., Lakehurst, 5 
May 1962, R. Hodges. 

alaskae Heinrich, 1923 (Epinotia cruci- 
ana): ] Bull. WW:S. Natl Muse 1239229. 
Holotype 6, USA, Alaska, Yukon, 3 
August 1916, G. Harrington [cited as 
*‘Huntington” by Heinrich]. The col- 
lecting locality, probably referring to 
Yukon River or some other locality in 
Alaska, is somewhat ambiguous _ be- 
cause Yukon Territory is in Canada, not 
Alaska. 

albafascia Heinrich, 1929 (Ancylis); Proc. 
U.S. Natl. Mus. 75: 19. Holotype ¢, 
USA, California, Tulare Co., Mineral 
King, “June 24-30.” 

albicapitana Busck, 1914 (Evetria); Proc. 
Entomol. Soc. Wash. 16: 147. Holotype 
6, Canada, Saskatchewan, Prince Al- 
bert, rf. Pinus divaricata, em: 12 March 
1914, J. Blumer. Although the type la- 
bel indicates ‘“‘Evetria capitana”’ rather 
than albicapitana, the USNM type 
number (Cat. No. 18444) and the col- 
lection data on the specimen match that 
given in the original description. 

albicaudana Busck, 1915 (Sparganothis); 
Proc. Entomol. Soc. Wash. 17: 85. Ho- 
lotype 36, USA, Pennsylvania, Notch, 
rf. maple, em: 5 July 1913, A. Busck. 

albiciliana Fernald, 1882 (Sericoris); Trans. 
Am. Entomol. Soc. 10: 70. Lectotype 
3d, USA, Maine, Penobscot Co., Orono. 
Designated by Miller (1970). 

albidula Heinrich, 1926 (Olethreutes buck- 
ellana); Bull. U.S. Natl. Mus. 132: 189. 
Holotype 6, USA, California, Inyo Co., 
15-30 June 1922, O. Poling. 

albimaculana Fernald, 1879 (Grapholitha); 
Canad. Entomol. 11: 157. Lectotype ¢, 


1018 


USA, Maine, Penobscot Co., Orono. 
Designated by Miller (1970). 

albipuncta Heinrich, 1926 (Bactra_ veru- 
tana); Bull. U.S. Natl. Mus. 132: 84. 
Holotype ¢, USA, Colorado, Adams 
Co., Denver, Oslar. 

albopunctana Brown, 1999 (Dimorphopal- 
pa); Pan-Pac. Entomol. 75: 85. Holo- 
type 6, Venezuela, Aragua, Rancho 
Grande, 1,100 m, 16—19 January 1966, 
S. & W. Duckworth. 

aliana Kawabe, 1965 (Clepsis); Kontyt 34: 
460. Holotype 6, Japan, Kurodake 
(Hokkaido), Mt. Daisetsu, 22 July 1952, 
A. Matsura. 

alishana Kawabe, 1986 (Epiblema); Ento- 
mol. Pap. Pres. Kurosawa, Tokyo: 82. 
Holotype 6, Taiwan, Chiai Hsien, Al- 
ishan, 2,300 m, 7 August 1971, Y. Shi- 
bata. 

alleniana Fernald, 1882 (Tortrix); Trans. 
Am. Entomol. Soc. 10: 68. “Type” dé, 
USA, Maine, Penobscot Co., Orono, 
July. This species was described from 
three males and three females; we have 
been able to locate only one male. The 
specimen is clearly labeled as “‘Type”’ 
in Fernald’s hand. 

allochroma Diakonoff, 1968 (Costosa); 
Bull. U.S. Natl. Mus. 257: 81. Holotype 
3, Philippine Islands, Davao Province, 
Mindanao, E slope Mt McKinley, 
5,600’, 16 September 1946, CNHM 
Philippine Zool. Exped. 1946—47, H. 
Hoogstraal. 

alphabetica Walsingham, 1914 (Eucosma); 
Biol. Centr.-Am., Lepid., Het. 4: 236. 
Holotype ¢, Mexico, Puebla, Popoca- 
tepetl [Park], 8—11,000’, July 1906, W. 
Schaus. Described from a single speci- 
men identified as “‘Type.”’ 

alphitopa Clarke, 1968 (Hysterosia); Proc. 
U.S. Natl. Mus. 125: 7. Holotype ¢, 
Venezuela, Aragua, Rancho Grande, 
16—23 October 1966, S. & W. Duck- 
worth. 

alterana Heinrich, 1923 (Thiodia); Bull. 
U.S. Natl. Mus. 123: 46. Holotype 6, 
USA, Maryland, Montgomery Co., 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Plummers Island, | August 1903, A. 
Busck. 

altissima Kawabe, 1978 (Eubrochoneura); 
Tinea 10: 177. Holotype 3, Japan, Hon- 
shu, Shizouka Pref., Odaru Spa, S Izu, 
29 April 1968, A. Kawabe. 

amatana Dyar, 1901 (Eulia); J. N. Y. En- 
tomol. Soc. 9: 24. Three syntypes (3 
22), USA, Florida, Palm Beach Co., 
Palm Beach, r.f. Nectandra [| = Ocotea| 
(n = 1). Dyar described this species 
from the three specimens cited above. 

amblyopa Clarke, 1976 (Cryptophlebia); 
Insects of Micronesia 9: 106. Holotype 
3, Micronesia, Palau Islands, Koror Is- 
land, Koror, January—May, light trap, 
Beardsley. 

ambodaidaleia Miller, 1983 (Phaneta); 
Ann. Entomol. Soc. Am. 76: 101. Ho- 
lotype 2, USA, South Carolina, Berke- 
ley Co., Wedge Plantation, Mc- 
Clellanville, 16 March 1968, D. Fergu- 
son. 

ambogonium Pogue, 1986 (Apolychrosis); 
in Cibridn-Tovar et al., Cone and Seed 
Insects of the Mexican Conifers: 21. 
Holotype ¢, Mexico, Mexico, San Mi- 
guel Atlautla, 2,500 m, rf. Pinus leio- 
phylla, em: 14 September 1983, W. San- 
chez. 

americana Fernald, 1882 (Teras); Trans. 
Am. Entomol. Soc. 10: 66. Lectotype 
3, USA, Massachusetts, ‘“‘“Cambr. B.”’ 
[Cambridge?]. Designated by Obraztsov 
(1963). 

amoena Kawabe, 1986 (Phaecasiophora); 
Entomol. Pap. Pres. Kurosawa, Tokyo: 
77. Holotype 6, Taiwan, Hualien Hsien, 
Hungyeh Spa, 200 m, 29-30 March 
1984, A. Kawabe. 

amorpha Clarke, 1986 (Dichelopa); Smith- 
sonian Contrib. Zool. 416: 123. Holo- 
type 6, Marquesas Islands, Nuku Hiva, 
Tapuaooa, 2,500’, 30 January 1968, J. 
Clarke. 

amphibola_ Diakonoff, 1982 (Diplosema- 
phora); Zool. Verhandel. (Leiden) 193: 
21. Holotype 6, Sn Lanka, Ratnapura 
District, Uggalkaltota, 350’, Irrigation 


VOLUME 102, NUMBER 4 


Bungalow, 31 January—8 February 
1970, D. Davis & B. Rowe. 

anaranjada Miller, 1959 (Laspeyresia); 
Florida Entomol. 42: 131. Holotype &, 
USA, Georgia, Crisp Co., Cordele, rf. 
Pinus ipalustris,,21 May %1950;) C. 
Speers. 

anastea Diakonoff, 1968 (Archidemis); 
Bull. U.S. Natl. Mus. 257: 30. Holotype 
36, Philippine Islands, Davao Province, 
Mindanao, E slope Mt McKinley, 
7,000’, 22 September 1946, CNHM 
Philippine Zool. Exped. 1946—47, H. 
Hoogstraal. 

anaxia Clarke, 1968 (Amallectis); Proc. 
U.S. Natl. Mus. 125: 32. Holotype °&, 
Guatemala, Volcan Santa Maria, July, 
Schaus and Barnes. 

andromedana Barnes & McDunnough, 
1917 (Olethreutes); Contrib. Nat. Hist. 
Lepid. North-Am. 3: 223. “Type” ¢, 
USA, Florida, Lee Co., Fort Meyers, r-f. 
Andromeda, “Apr 24-30.’ According 
to the original description, this species 
was described from two males and four 
females. Although a holotype was not 
designated, the specimen cited above is 
clearly labeled “Type.” 

angusana Fernald, 1892 (Peronea); Canad. 
Entomol. 24: 178. Two syntypes (1 <4, 
1 2), USA, New York, West Farms, J. 
Angus; and USA, Maine, Penobscot 
Co., Orono, 1 August 1884. According 
to the original description these two 
specimens comprise the type series. 

animosana Busck, 1907 (Tortrix); J. N. Y. 
Entomol. Soc. 15: 235. Holotype °&, 
Mexico, Jalapa, W. Schaus. 

anisoneura Diakonoff, 1982 (Eupoecilia); 
Zool. Verhandel. (Leiden) 193: 6. Ho- 
lotype ¢, Sri Lanka, Ratnapura District, 
Uggalkaltota, 350’, Irrigation Bunga- 
low, 31 January—8 February 1970, D. 
Davis & B. Rowe. 

anisoptera Clarke, 1976 (Dudua); Insects 
of Micronesia 9: 95. Holotype 6, Mi- 
cronesia, Guam, Mt. Chachao, 16 May 
1936, O. Swezey. 


anthrocodelta Clarke, 1971 (Dichelopa); 


1019 


Smithsonian Contrib. Zool. 56: 104. 
Holotype 3, Rapa Island, Maugaoa, 
950’ (292 m), 23 November 1963, J. & 
T. Clarke. 

aperta Diakonoff, 1968 (Archips); Bull. 
U.S. Natl. Mus. 257: 28. Holotype 2, 
Philippine Islands, Luzon, Mountain 
Province, Baguio, 7 May 1945, J. Fran- 
clemont. 

aporema Dognin, 1912 (Cnephasia); Het- 
eroceres Nouveaux de LAmerique du 
Sud 6: 49. Holotype ¢, Colombia, Fas- 
sel. 

aporrhegma Clarke, 1986 (Dichelopa phal- 
aranthes); Smithsonian Contrib. Zool. 
416: 136. Holotype 6, Marquesas Is- 
lands, Nuku Hiva, Tapuaooa, 2,500’, 30 
January 1968, J. & T. Clarke. 

apospasta Obraztsov, 1964 (Proeulia); 
Proc. U.S. Natl. Mus. 116: 191. Holo- 
type @, Chile, Concepcion, October 
1902, E. Reed. 

approximana Heinrich, 1919 (Olethreutes); 
Insect. Inscit. Menst. 7: 65. Holotype 3, 
USA, New York, Rensselaer, rolling ter- 
minal leaves of loosestrife [Lysima- 
chiala12 June 19116, NYS Coll: 

aprilana Grote, 1877 (Exentera); Canad. 
Entomol. 9: 227. Two syntypes (1 d, 1 
2), USA, New York, Albany Co., Al- 
bany, April. According to the original 
description, this species was described 
from one male and one female collected 
the ‘‘end of April” by Lintner and Hill. 
The syntypes consist only of wings 
glued to pieces of cardboard: two fore- 
wings of the male, one forewing of the 
female. The sexes of the specimens are 
indicated on the specimen labels. 

aquila Busck, 1914 (Homona); Proc. U.S. 
Natl. Mus. 47: 53. Holotype 6, Panama, 
Cabima, May 1911, A. Busck. 

arammclaina Razowski, 1987 (Heppnero- 
grapha); Bull. Polish Acad. Sci. 35: 64. 
Holotype ¢, Costa Rica, Cartago Prov- 
ince, Tuis, 28 May—4 June, W. Schaus. 

argema Clarke, 1986 (Dichelopa); Smith- 
sonian Contrib. Zool. 416: 152. Holo- 
type 2, Marquesas Islands, Fatu Hiva, 


1020 


Mt. Teoaiua, 2,000’, 22 March 1968, J. 
& T. Clarke. 

argentifasciata Heppner, 1989 (Choristo- 
neura); Florida Entomol. 72: 104. Ho- 
lotype 6, USA, Florida, Glades Co., 
Fisheating Creek, Palmdale, 7-10 May 
1964, R. Hodges. 

argentifurcatana Grote, 1876 (Conchylis); 
Canad. Entomol. 8: 206. Syntype ¢é, 
Canada, Ontario, Port Stanley, W. Saun- 
ders. Although the specimen bears a la- 
bel “‘London, Ont.,” according to the 
original description the specimens were 
taken at Port Stanley, by ““W. Saunders 
from London, Ontario.”’ The disposition 
of the other syntype is unknown to us. 

argentina Brown, 1991 (Cuproxena); in 
Brown and Powell, Univ. Calif. Publ. 
Entomol. 111: 64. Holotype 2, Argen- 
tina, Misiones, Puerto Rico, 4—8 April 
LOW, Crs2O, Flint. 

argillacea Clarke, 1976 (Ruthita); Insects 
of Micronesia 9: 26. Holotype 2, Mi- 
cronesia, Guam, May 1936, O. Swezey. 

argodonta Clarke, 1955 (Orthocomotis); 
Trans. Royal Entomol. Soc. London 
107g lS4 yHoletype 925) Brazile St 
Catherines”’ [Santa Catarina], EK John- 
son. 

argutipunctana Blanchard & Knudson, 
1983 (Phaneta); J. Lepid. Soc. 37: 143. 
Holotype ¢d, USA, Texas, Hemphill 
Co., Canadian, 15 August 1971, A. & 
M. E. Blanchard. 

argyraspis Razowski, 1984 (Saphenista); 
Ann. Zool. Warsaw 38: 278. Holotype 
2, Venezuela, Distrito Federal, 14 km 
NE Tovar, 28—29 February 1976, C. & 
O. Flint. 

argyrospiloides Clarke, 1971 (Dichelopa); 
Smithsonian Contrib. Zool. 56: 118. 
Holotype 2, Rapa Island, Maurua, 600’ 
(184 m), 25 October 1963, J. Clarke. 

arizonana Powell, 1975 (Epiblema); Pan- 
Pac. Entomol. 51: 101. Holotype ¢, 
USA, Arizona, Coconino Co., Oak 
Creek Canyon, Flagstaff, rf. Rudbeckia 
laciniata, em: 28 June 1936, G. Engle- 
hardt. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


arizonensis Heinrich, 1914 (Evetria albi- 
capitana); Proc. U.S. Natl. Mus. 57: 57. 
Holotype 6, USA, Arizona, Santa Cat- 
alina Mountains, r.f. Pinus cembroides, 
23 June 1917, Hopkins no. 13977, G. 
Hofer. 

arrhostia Clarke, 1968 (Phtheochroa); 
Proc. U.S. Natl. Mus. 125: 56. Holotype 
2, Peru, Cusco, Machu Picchu, 2,385 
m, 5 March 1959, J. Clarke. 

arthuri Dang, 1984 (Cochylis); Canad. En- 
tomol. 116: 253. Holotype ¢, Canada, 
Saskatchewan, S.E. Saskatoon, r.f. He- 
lianthus sp., August 1980. 

asthenia Clarke, 1968 (Carolella); Proc. 
U.S. Natl. Mus. 125: 55. Holotype 4, 
Guatemala, Palin, July, Schaus & 
Barnes. 

asynthetes Diakonoff, 1968 (Gatesclark- 
eana); Bull. U.S. Natl. Mus. 257: 42. 
Holotype 6, Philippine Islands, Cadiz, 
Occ. Negros, **3/8/29,”’ Batabas. 

atascosana Blanchard, 1979 (Eucosma); J. 
Lepid. Soc. 33: 212. Holotype ¢, USA, 
Texas, Cameron Co., Laguna Atascosa, 
22 November 1973, A. & M. Blan- 
chard. 

atayalicana Kawabe, 1989 (Acleris); Tinea 
12: 195. Holotype 3, Taiwan, Hualien 
Hsien, Tayuling, forest, 2,500 m, 9-18 
June 1980, D. Davis. 

atomosana Busck, 1907 (Phalonia); J. N. 
Y. Entomol. Soc. 15: 22. Holotype 6, 
USA, Pennsylvania, Allegheny Co., 
Pittsburgh, 17 August 1905, H. Engel. 

atomosana Walsingham, 1879 (Paedisca); 
Ill. Lepid. Heter. Brit. Mus. 4: 42. Syn- 
type ¢, USA, California [San Francisco 
or Bear Valley according to the original 
description], May—June, 1871. This spe- 
cies was described from two males; the 
other syntype is presumably in the 
BMNH. 

atrata Diakonoff, 1973 (Lobesia); Zool. 
Monogr. Rijksmus. Nat. Hist. 1: 382. 
Holotype ¢, Marianas Islands, Guam, 
Piti, “‘7/9/36,”” O. Swezey. 

atrilinea Clarke, 1976 (Cryptophlebia); In- 
sects of Micronesia 9: 115. Holotype °, 


VOLUME 102, NUMBER 4 


Micronesia, Ponape, Colonia, 14 Janu- 
ary 1953, J. Clarke. 

atriplaga Clarke, 1976 (Duessa); Insects of 
Micronesia 9: 28. Holotype 2, Micro- 
nesia, Rota, nr. Sabana, 21 June 1946, 
Townes 807, at light. 

atristriga Clarke, 1953 (Epinotia); J. Wash. 
Acad. Sci. 43: 228. Holotype 6, USA, 
Illinois, Putnam Co., 24 March 1938, 
M. Glenn. Although the collection date 
of the holotype is cited as “‘March 17, 
1945”’ in the original description, the 
specimen listed above is unambiguously 
labeled as the holotype. 

atrodentana Fernald, 1882 (Eccopsis); 
Trans. Am. Entomol. Soc. 10: 71. Lec- 
totype ¢, Canada, Ontario, London, W. 
Saunders. Designated by Miller (1970). 

atsushii Bae, 1993 (Lobesia); Japan. J. En- 
tomol. 61: 516. Holotype 6, Taiwan, 
Hualien Hsien, Mt. Hohuanshan, 3,100 
m, 30 July—1 August 1983, A. Kawabe. 

audaculana Busck, 1907 (Archips); J. N. Y. 
Entomol. Soc. 15: 235. Holotype °, 
Mexico, Veracruz, Orizaba, R. Muller. 

aurantia Clarke, 1976 (Adoxophyes); In- 
sects of Micronesia 9: 127. Holotype 2, 
Micronesia, Ponape, Colonia, 12—17 
January 1953, J. Clarke. 

aurantica Busck, 1920 (Epagoge); Insect. 
Inscit. Menst. 8: 84. Holotype 2, Costa 
Rica, Cartago Province, Juan Vinas, W. 
Schaus. 

auraria Clarke, 1949 (Eulia); Acta Zool. 
Lilloana (Tucuman) 7: 583. Holotype 
3, Chile, Santiago Province, Cajon de 
Maypo, Cordillera, El Cencio, 12—20 
January 1948, T. Ramirez. 

aurata Diakonoff, 1968 (Adoxophyes); 
Bull. U.S. Natl. Mus. 257: 11. Holotype 
3, Philippine Islands, Luzon, Mt. Mak- 
iling, Baker. 

aureana Busck, 1907 (Phalonia); J. N. Y. 
Entomol. Soc. 15: 26. Holotype 9°, 
USA. Pennsylvania, Allegheny Co., 
Oak Station, 17 June 1906, FE Marloff. 

aureola Diakonoff, 1977 (Nexosa); Zool. 
Verhandel. (Leiden) 158: 15. Holotype 
2, British New Guinea, Hydrographer 


1021 


Mountains, 2,500’, March 1918, Ei- 
chhorn Bros. 

auricaput Razowski, 1971 (Acleris); Acta 
Zool. Cracov. 16: 550. Holotype °, Tai- 
wan, Hassenzan, 6 June 1942, S. Issik1i. 

aurichalceana Riley, 1881 (Melissopus); 
Trans) StieouishAcad: «Seis 41323: 
“Type” ¢, USA, Fernald, rf. acorns, 
**1/6/76.”” Riley did not mention the 
number of specimens he examined; this 
is the only Riley specimen of aurichal- 
ceana we have been able to locate. It is 
labeled *““Type”’ in Riley’s hand. 

auricomana Busck, 1907 (Tortrix); J. N. Y. 
Entomol. Soc. 15: 236. Holotype 6, 
Mexico, June 1906, R. Muller. 

auriferana Busck, 1911 (Tortrix); Proc. 
U.S. Natl. Mus. 40: 227. Holotype ¢, 
Brazil, Parana, Castro, W. Schaus, 
“6187 Wlsm. 1908.” 

austrina Miller, 1985 (Eucosma); Ann. En- 
tomol. Soc. Am. 78: 243. Holotype ¢, 
USA, Texas, Hemphill Co., Canadian, 
15 August 1971, A. & M. Blanchard. 

azumina Yasuda & Kawabe, 1980 (Ar- 
chips); Tinea 11: 13. Holotype @, Ja- 
pan, Nagano Pref., Azumi V., Shimaji- 
ma-Dan, 780 m, 15 April 1977, N. Hi- 
bano. 


baccharivora Pogue, 1988 (Lorita); Proc. 
Entomol. Soc. Wash. 90: 449. Holotype 
6, USA, Florida, Okeechobee Co., 
Lake Okeechobee, 27°N, 81°W, Septem- 
ber 1984 (lab. reared), D. Green. 

bactrana Heinrich, 1923 (Eucosma); Bull. 
U.S. Natl. Mus. 123: 117. Holotype ¢, 
USA, Colorado, San Juan Co., Silver- 
ton, “July 16—23.” 

baea Razowski, 1987 (Transtillaspis); Bull. 
Polish Acad. Sci. 35: 79. Holotype <6, 
Colombia, Narino, Volcan Galeras, 
2,900 m, 13 January 1959, J. Clarke. 

bakeri Diakonoff, 1968 (Homona); Bull. 
U.S. Natl. Mus. 257: 20. Holotype ¢, 
Philippine Islands, Luzon, Mt. Makil- 
ing, Baker. 

balia Diakonoff, 1982 (Acanthoclita); Zool. 
Verhandel. (Leiden) 193: 30. Holotype 


1022 


6, Sri Lanka, Ratnapura District, Ug- 
galkaltota, 350’, Irrigation Bungalow, 
31 January—8 February 1970, D. Davis 
& B. Rowe. 

balioleuca Clarke, 1976 (Adoxophyes); In- 
sects of Micronesia 9: 136. Holotype 6, 
Micronesia, Ponape, N. slope Tamata- 
mansakir, 19 January 1953, J. Clarke. 

bana Kearfott, 1907 (Phalonia); Trans. Am. 
Entomol: “Soci9332775) Lectotype: d, 
USA, Illinois, Cook Co., Chicago, June, 
J. Reading. Designated by Klots (1942). 

banana Busck, 1906 (Lipoptycha); Proc. 
Biol. Soc. Wash. 19: 182. Holotype ¢, 
USA, Colorado, South Park, Oslar. 

baracana Busck, 1907 (Aysterosia); J. N. 
Y. Entomol. Soc. 15: 33. Holotype 6, 
USA, Missouri, St. Louis Co., St. Louis, 
11 July 1906, McElhose. 

bascanion Razowski, 1987 (Transtillaspis); 
Bull. Polish Acad. Sci. 35: 75. Holotype 
6, Peru, Cusco, Machu Picchu, 2,700 
m, 6 February 1959, J. Clarke. 

basipunctana Walsingham, 1879 (Paedis- 
ca); Ill. Lepid. Heter. Brit. Mus. 4: 40. 
Syntype 2, USA, California, ‘“‘Lower 
Lake,”’ 22 June 1871. Walsingham de- 
scribed this species from two males and 
three females; the other syntypes are 
presumed to be in the BMNH. 

batesi Heinrich, 1932 (Talponia); Proc. En- 
tomol. Soc. Wash. 34: 20. Holotype 6, 
Guatemala, Antiqua, 5,000’, rf. Annona 
cherimoia, M. Bates. 

batoidea Razowski, 1987 (Transtillaspis); 
Bull. Polish Acad. Sci. 35: 75. Holotype 
3, Peru, Cusco, Machu Picchu, 5 Feb- 
ruary 1959, J. Clarke. 

bauhiniae Busck, 1934 (Ancylis); Entomol. 
Am. 13: 155. Holotype 6, Cuba, San- 
tiago de Vegas, Hab., leaftier in Bauhi- 
nia heterophylla, 4 December 1931, R. 
Olera. 

bebela Razowski, 1987 (Transtillaspis); 
Bull. Polish Acad. Sci. 35: 82. Holotype 
3d, Colombia, Bogota, Chico, 25 Janu- 
ary 1959) ad#Clarke: 

beckeri Clarke, 1973 (Eumarozia); J. Lepid. 
Soc. 27: 269. Holotype 6, Costa Rica, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Cartago Province, Turrialba, rf. Jug- 
lans, em: 29 August 1972, V. Becker. 

benjamini Heinrich, 1923 (Thiodia); Bull. 
U.S. Nat. Mus. 123: 66. Holotype ¢, 
USA, Utah, Vineyard, 12 September 
1912, T. Spalding. 

bicolor Kawabe, 1963 (Acleris); Tyo to Ga 
14: 70. Holotype 6, Japan, Nagano 
Pref., Honshu, Naka-karuisawa, 23 Sep- 
tember 1962, T. Maenami. 

bicolor Kawabe, 1978 (Aterpia); Tinea 10: 
174. Holotype ¢, Japan, Tsushima Is- 
land, Taterayama, 2 November 1973, T. 
Watanabe. 

bicolor Powell, 1961 (Decodes); J. Lepid. 
Soc. 14: 122. Holotype 2, USA, Cali- 
fornia, Napa Co., Mt. St. Helena, 18 
April 1939, E. Johnston. 

bicolora Kawabe, 1976 (Zeiraphera); Tinea 
10: 42. Holotype ¢, Japan, Gunma 
Pref., Kumanotaira, 14 July 1952, M. 
Hoshino. 

bicornigera Razowski, 1984 (Histura); 
Acta Zool. Cracov. 27: 212. Holotype 
3, Colombia, Fassel. 

bicornis Diakonoff, 1968 (Homona); Bull. 
U.S. Natl. Mus. 257: 18. Holotype 6, 
Philippine Islands, Luzon, Los Banos, 
Baker. 

biemina Kawabe, 1980 (Apotomis); Tinea 
11: 18. Holotype ¢, Japan, Miyagi 
Pref., Tokatta, 13 September 1971, W. 
Watanabe. 

bigaulae Brown, 1999 (Eubetia); J. New 
York Entomol. Soc. 106: 181. Holotype 
6, Venezuela, Aragua, Rancho Grande, 
cloud forest, 1,100 m, 30—31 March 
1978, blacklight, J. Heppner. 

binotata Brown & Obraztsov, 1991 (Cu- 
proxena); in Brown and Powell, Univ. 
Calif. Publ. Entomol. 111: 61. Holotype 
?, Brazil, Santa Catarina, 22 July 1935, 
FE Hoffman. 

biquadrana Walsingham, 1876 (Paedisca); 
Ill. Lepid. Heter. Brit. Mus. 4: 45. Syn- 
type 6, USA, California, Shasta Co., 
Pitt River, “end of July” 1871. Wal- 
singham described this species from 


VOLUME 102, NUMBER 4 


two males; the other syntype is pre- 
sumed to be in the BMNH. 

bira Kawabe, 1976 (Endothenia); Tinea 10: 
43. Holotype 6, Japan, Ishikawa Pref., 
Anamizu, Bira, 18 August 1961, A. Ka- 
wabe. 

birdana Busck, 1907 (Hysterosia); J. N. Y. 
Entomol. Soc 15: 32. Holotype 9°, 
USA, New York, Rye, “‘bores Helian- 
thus,’ H. Bird. 

biserrata Brown, 1991 (Paraptila); J. Lep- 
id. Soc. 44: 269. Holotype ¢, Costa 
Rica, Cartago Province, Turrialba, 22— 
28 February 1965, S. & W. Duckworth. 

bittana Busck, 1906 (Hemimene); Proc. 
Biol. Soc. Wash. 19: 179. Holotype ¢, 
USA, Pennsylvania, Allegheny Co., 
Pittsburgh, 29 May 1905, H. Engel. 

blackmorei Obraztsov, 1963 (Acleris emar- 
gana); Proc. U.S. Natl. Mus. 114: 208. 
Holotype ¢, Canada, British Columbia, 
Goldstream, 7 August 1923, E. Black- 
more. 

blanchardi Miller, 1978 (Rhyacionia); in 
Powell and Miller, U.S. Dept. Agric., 
Agric. Handb. 514: 19. Holotype ¢d, 
USA, Texas, Montgomery Co., Conroe, 
10 March 1968, A. & M. Blanchard. 

blechra Razowski, 1987 (Transtillaspis); 
Bull. Polish Acad. Sci. 35: 79. Holotype 
3, Colombia, Narifio, Volcan Galeras, 
2,900 m, 13 January 1959, J. Clarke. 

bobana Kearfott, 1907 (Eucosma); Trans. 
Am. Entomol. Soc. 33: 26. ‘“‘Type’’ 3, 
USA, Colorado, Salida, 11 June 1888, 
W. Dietz. This species was described 
from three ‘“‘co-types”’ (Salida, Colora- 
do; Southwestern Colorado; Harris Co., 
Texas). Although Heinrich (1923) indi- 
cated that the “type” (from Salida, Col- 
orado) was in the AMNH, Klots (1942) 
correctly identified the USNM as the 
place of disposition. Klots designated 
the AMNH specimen (Southwestern 
Colorado) as a “‘lectoparatype.”” The 
disposition of the third syntype is un- 
known. 

bolanderana Walsingham, 1876 (Paedis- 
ca); Ill. Lepid. Heter. Brit. Mus. 4: 42. 


1023 


Two syntypes (2 66), USA, California, 
Siskiyou Co., Mount Shasta, August 
1871. This species was described from 
three males and two females; the other 
syntypes are presumed to be in the 
BMNH. 

boliviae Razowski, 1988 (Proeulia); Acta 
Zool. Cracov. 31: 407. Holotype 6, Bo- 
livia, Cochabamba, Incachaca, tropical 
cloud area, 2,100 m, 27 August—5 Sep- 
tember 1956, L. Pefia. 

boliviana Brown, 1991 (Bidorpitia); in 
Brown and Powell, Univ. Calif. Publ. 
Entomol. 111: 75. Holotype @, Bolivia, 
Cochabamba, Incachaca, tropical cloud 
area, 2,100 m, 27 August—5 September 
1956, Lk. Pena: 

boliviana Razowski, 1984 (Histura); Acta 
Zool. Cracov. 27: 212. Holotype 3, Bo- 
livia, Cochabamba, Incachaca, tropical 
cloud area, 2,100 m, 27 August—5 Sep- 
tember 1956, L. Pefia. 

boscantica Dognin, 1912 (Tortrix); Heter- 
oceres L-Amerique du Sud 6: 48. Ho- 
lotype ¢, Colombia, Cali, San Antonio, 
Fassel. 

bourquini Clarke, 1949 (Eulia); Acta Zool. 
Lilloana (Tucuman) 7: 581. Holotype 
?, Brazil, Parana, Castro, W. Schaus. 

brachistocera Razowski, 1987 (Transtillas- 
pis); Bull. Polish Acad. Sci. 35: 81. Ho- 
lotype 6, Colombia, Bogota, Chico, 25 
January 1959, J. Clarke. 

brachystigma Clarke, 1965 (Nesochoris); 
Proc. U.S. Natl. Mus. 117: 76. Holotype 
3d, Chile, Juan Fernandez Island, Mas- 
atierra, E] Rabanal, 350 m, 27 February 
195i 0P) Kuschel: 

bracteatana Fernald, 1881 (Grapholitha); 
in Comstock, Annu. Rept. Dept. Agric. 
1881: 265. Lectotype 6, USA, Califor- 
nia, Jolon, em: 13 September 1880. 
Designated by Miller (1970). 

bramiliana Brown, 1991 (Cuproxena); in 
Brown and Powell, Univ. Calif. Publ. 
Entomol. 111: 61. Holotype ¢, Brazil, 
Rio de Janeiro, Petropolis, 600 m, 10 
October 1985, S. Miller. 

brandinojuxta Razowski, 1987 (Transtillas- 


1024 


pis); Bull. Polish Acad. Sci. 35: 77. Ho- 
lotype ¢, Bolivia, Cochabamba, Inca- 
chaca, tropical cloud area, 2,100 m, 27 
August—5 September 1956, L. Pena. 

brauni Heinrich, 1931 (Anchylopera); Proc. 
U.S. Natl. Mus. 79: 11. Holotype 4, 
USA, Ohio, Adams Co., Beaver Pond, 
11 May 1927. 

braziliana Brown, 1991 (Punctapinella); 
Los Angeles Co. Mus. Contrib. Zool. 
423: 4. Holotype °, Brazil, Santa Ca- 
tarina, 20 November 1936, E Hoffman. 

britana Busck, 1906 (Hemimene); Proc. 
Biol. Soc. Wash. 19: 178. Lectotype ¢, 
Canada, British Columbia, Kaslo, H. 
Dyar. Designated by Miller (1983). 

britannia Kearfott, 1904 (Acleris); Canad. 
Entomol. 36: 138. Holotype ¢, Canada, 
British Columbia, Kaslo, H. Dyar. 

broui Knudson, 1986 (Dichrorampha); J. 
Lepid. Soc. 40: 323. Holotype 6, USA, 
Louisiana, St. Tammany Parish, 4.2 mi 
NE Abita Springs, sec. 24, T6, SR 128, 
uv [light], 12 April 1985,V. Brou. 

brunana Brown, 1990 (Hynhamia); Ento- 
mol. Scand. 21: 325. Holotype d, Peru, 
Angasmarca. 

brunneopurpuratum Heinrich, 1923 (Exar- 
tema); Proc. Entomol. Soc. Wash. 25: 
118. Holotype 2, USA, Virginia, Fair- 
fax Co., Falls Church, rf. alder, em: 1 
August 1913. 

bumeliana Heinrich, 1926 (Goditha); Bull. 
U.S. Natl. Mus. 132: 8. Holotype 6, 
USA, Texas, Dallas Co., Dallas, “‘Di- 
chrorampha boumelliana Boll mss., 
two generations, June and Oct. in the 
rolled leaves of Boumellia languinosa.” 
Although misspelled as ‘‘rumeliana” 
immediately prior to its description, it is 
spelled correctly as “‘bumeliana’’ pre- 
viously on the same page, where it is 
identified as the type species of Godi- 
tha. 

burgessiana Zeller, 1875 (Phoxopteris); 
Verhandel. Zool.-Bot. Ges. Wein 25: 
252. “Type” ¢°, USA, Massachusetts, 
Essex Co., Beverly, 18 June 1869. This 
species was described from a male and 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


two females; the specimen listed above 
is the only Zeller specimen of burges- 
siana we could find. 

bununa Kawabe, 1989 (Acleris); Tinea 12: 
196. Holotype 2, Taiwan, Hualien 
Hsien, Tayuling, 2,560 m, 1 April 1984, 
A. Kawabe. 

busckana J. A. Comstock, 1939 (Pharma- 
cis); Bull. South. Calif. Acad. Sci. 38: 
112. Holotype ¢, USA, California, Los 
Angeles Co., El Segundo, em: 16 No- 
vember 1938, W. Pierce. 

busckana Heinrich, 1923 (Rhyacionia); 
Bull. U.S. Natl. Mus. 123: 17. Holotype 
3d, USA, New York, Nassau Co., Long 
Island, Bellmore, April 1913, G. Engle- 
hardt. 

busckana Keifer, 1933 (Clepsis); Calif. 
Dept. Agric. Mon. Bull. 22: 351. Ho- 
lotype 6, USA, California, San Francis- 
co, 8 February 1933, rf. Scophularia 
californica, em: 3 March 1933, H. Kei- 
fer. 

buscki Brown, 2000 (Accuminulia); J. Lep- 
id. Soc. 53: 61. Holotype 3, Chile, San- 
tiago Province, reared from grape 
(fruit), em: 12 April 1954, M. Ramsay. 

bushiensis Kawabe, 1980 (Epinotia); Tinea 
11: 22. Holotype 6, Japan, Saitama 
Pref., Bushi, Iruma, 20 October 1978, 
H. Inoue. 

bushnelli Busck, 1914 (Evetria); Proc. En- 
tomol. Soc. Wash. 66: 144. Holotype @, 
USA, New Mexico, Ft. Bayard, rf. Pi- 
nus ponderosa, em: 19 March 1914, C. 
Bushnell. 


caeruleana Walsingham, 1879 (Grapholi- 
tha); Ill. Lepid. Heter. Brit. Mus. 4: 66. 
Syntype 6, USA, Oregon, Rouge River, 
May 1872. Walsingham described this 
species from one male and one female; 
the female is presumed to be in the 
BMNH. 

caeruleumana Kawabe, 1980 (Zeiraphera); 
Tinea 11: 24. Holotype 3, Japan, Akita 
Pref., Tamagawa, 2 September 1970, T. 
Watanabe. 


caesiata Clarke, 1968 (Cochylis); Proc. 


VOLUME 102, NUMBER 4 


U.S. Natl. Mus. 125: 18. Holotype °&, 
Venezuela, Aragua, Rancho Grande, 
1,100 m, 16—23 October 1966, S. & W. 
Duckworth. 

californiae Heinrich, 1923 (Hystricophora 
strygiana); Bull. U.S. Natl. Mus. 123: 
256. Holotype ¢, USA, California, 
Lake Tahoe, Deer Park Springs, “‘July 
8—15.’’ Heinrich (1923) consistently 
misspelled Hystrichophora in his revi- 
sion. 

callosoma Clarke, 1976 (Cryptophlebia); 
Insects of Micronesia 9: 112. Holotype 
2, Micronesia, Guam, Pt. Oca, near 
Agana, 12 May 1945, Bohart & Gres- 
sitt. 

campestrana Zeller, 1875 (Sericoris); Ver- 
handel. Zool.-Bot. Ges. Wein 25: 282. 
“Type” 6, USA [Maine or Massachu- 
setts according to the original descrip- 
tion], Packard. Zeller did not mention 
the number of specimens examined, al- 
though it is clear that he had both sexes. 
This is the only Zeller specimen of cam- 
pestrana that we were able to find. 

campicolana Walsingham, 1879 (Cochylis); 
Ill. Lepid. Heter. Brit. Mus. 4: 29. Two 
syntypes (6, 2), USA, California, Men- 
docino Co., 10 June 1871. Walsingham 
described this species from three males 
and two females; the other syntypes are 
presumed to be in the BMNH. 

canariana Barnes & Busck, 1920 (Ayster- 
osia); Contrib. Nat. Hist. Lepid. North 
Am. 4: 218. Holotype 6, USA, Arizo- 
na, White Mountains. 

candidus Pogue, 1986 (Apolychrosis); in 
Cibrian-Tovar et al., Cone and Seed In- 
sects of the Mexican Conifers: 23. Ho- 
lotype 6, Mexico, Puebla, Atotocoyan, 
rf. Pinus strobus, em: 2 October 1981, 
T. Mendez. 

canitia Clarke, 1986 (Dichelopa); Smith- 
sonian Contrib. Zool. 416: 149. Holo- 
type °, Marquesas Islands, Nuku Hiva, 
Tapuaooa, 2,500’, 30 January 1968, J. 
& T. Clarke. 

canofascia Forbes, 1930 (Olethreutes); Sci. 
Surv. Porto Rico and Virgin Islands 12: 


1025 


86. Holotype 3, Puerto Rico, Rio Pie- 
dras, 10 July 1916, E. Smith. 

canusana Wright, 1997 (Phaneta); in 
Wright, Brown and Gibson, J. Lepid. 
Soc. 51: 122. Holotype ¢, USA, Ohio, 
Adams Co., Lynx Prairie Preserve, Sta- 
tion 6, 17 March 1989, D. Wright. 

capitana Busck, 1906 (Hemimene); Proc. 
Biol. Soc. Wash. 19: 178. Holotype 3, 
USA, Colorado, South Park, Oslar. 

capizziana Obraztsov, 1963 (Acleris); Proc. 
U.S. Natl. Mus. 114: 252. Holotype °, 
USA, Oregon, Bendon, 24 September 
1956, J. Capizzi. 

capronata Razowski, 1988 (Ernocornutia); 
Acta Zool. Cracov. 31: 398. Holotype 
6, Colombia, Cauca, Paramo de Parace, 
Lake San Rafael, 3,570 m, 27 January 
195934).. Clarke: 

carduana Busck, 1907 (Polychrosis); J. N. 
Y. Entomol. Soc. 15: 134. Holotype &, 
USA, Maryland, Montgomery Co., Hy- 
attsville, rf. thistle, August 1906, A. 
Busck. 

carnana Barnes & Busck, 1920 (Tortrix); 
Contrib. Nat. Hist. Lepid. North Am. 4: 
214. Holotype 6, USA, California, San 
Bernardino Co., San Bernardino Moun- 
tains, Camp Baldy, “‘June 24—20.” 

carpophagoides Clarke, 1951 (Cryptophle- 
bia); J. Wash. Acad. Sci. 41: 299. Ho- 
lotype 6, Argentina, Tucuman, rf. seeds 
of pacara, K. Hayward. 

caryana Fitch, 1856 (Grapholitha); New 
York Agric. Rept. 16: 459. Two syn- 
types (2 6d), USA, New York [no data 
on specimen labels],“‘14,957,’’ Type 
No. 394. Fitch did not indicate the num- 
ber of specimens examined. 

cataclasta Diakonoff, 1982 (Dicnecidia); 
Zool. Verhandel. (Leiden) 193: 42. Ho- 
lotype ¢, Sri Lanka, Anuradhapura Dis- 
trict, Wildlife Sanctuary Bungalow, Hu- 
nuwipagama, Wilpattu, 200’, 10-19 
March 1970, D. Davis & B. Rowe. 

cathedra Clarke, 1976 (Lobesia); Insects of 
Micronesia 9: 105. Holotype 6, Micro- 
nesia, Guam, Pt. Oca, 30 May 1945, 
Bohart & Gressitt. 


1026 


catopta Razowski, 1988 (Ernocornutia); 
Acta Zool. Cracov. 31: 397. Holotype 
6, Colombia, Cauca, Paramo de Parace, 
Lake San Rafael, 3,570 m, 27 January 
1959, J: Clarke. 

caulocatax Razowski, 1984 (Conchylis); 
Ann. Zool. Cracov. 38: 278. Holotype 
36, Venezuela, Bolivar, Morichal Tauca, 
22 km E Rio Caura, 8—9 February 1976, 
C. & O. Flint. 

celiae Clarke, 1976 (Laspeyresia); Insects 
of Micronesia 9: 120. Holotype 2, Mi- 
cronesia, Yap Islands, Yap Island, Co- 
lonia, 21 June 1957, C. Sabrosky. 

celtisana Riley, 1881 (Paedisca); Trans. St. 
Louis Acad. Sci 4: 319. Holotype ¢, 
USA, Texas, rf. Celtis, Boll. Riley de- 
scribed this species from one specimen. 

cephalanthana Heinrich, 1921 (Phalonia); 
J. Agric. Res. 20: 825. Holotype ¢, 
USA, Texas, Charlotte, rf. Cephalan- 
thus occidentalis, em: 16 September 
1918, C. Heinrich. 

cercocarpana Dyar, 1903 (Eucosma); Proc. 
Entomol. Soc. Wash. 5: 297. Three syn- 
types (3 66), USA, Colorado, Platte 
Canyon, r.f. Cercocarpus parvifolius, 
Dyar & Caudell. Dyar described this 
species from three males and one fe- 
male. 

cerinus Kawabe, 1978 (Neoansthamna); 
Tinea 10: 182. Holotype 6, Japan, Hon- 
shu, Nippara, Tokyo, 20 June 1964, A. 
Kawabe. 

cervinana Fernald, 1882 (Teras); Trans. 
Am. Entomol. Soc. 10: 65. Lectotype 
3d, USA, Massachusetts, Cambridge (7). 
Designated by Obraztsov (1963). 

chalcana Packard, 1867 (Conchylis); Proc. 
Boston Soc. Nat. Hist. 11: 56. ““Type”’ 
(sex unknown; only a forewing), Can- 
ada, Labrador, Strawberry Harbor, near 
Cape Webuc, 26 July. The description 
does not indicate how many specimens 
Packard examined, but the species is 
listed as “‘uncommon.”’ 

chalybeana Fernald, 1882 (Peronea); 
Trans. Am. Entomol. Soc. 10: 65. Two 
syntypes (1 ¢6, 1 2), USA, New York, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


“L.I.”’ [Long Island]. Of the three spec- 
imens referred to by Fernald (1.e., two 
males and one female from Orono, 
Maine and New York), only the two 
specimens from New York have been 
located by us. 

changi Kawabe, 1989 (Capua); Tinea 12: 
192. Holotype 6, Taiwan, Haulien 
Hsien, Hohuanshan, 3,100 m, 30 July— 
1 August 1983, A. Kawabe. 

charma Clarke, 1968 (Phalonia); Proc. 
U.S. Natl. Mus. 125: 16. Holotype 2°, 
Argentina, Tucuman, Ciudad Universi- 
taria, 800 m, 17 February 1959, J. 
Clarke. 

chica Brown, 1984 (Corticivora); Proc. En- 
tomol. Soc. Wash. 86: 283. Holotype 3, 
USA, Florida, Highlands Co., Lake 
Placid, Archbold Biol. Sta., 1-7 May 
1964, R. Hodges. 

chiquitana Barnes & Busck, 1920 (Platy- 
nota); Contrib. Nat. Hist. Lepid. North 
Am. 4: 213. Holotype 6, USA, Califor- 
nia, San Bernardino Co., Loma Linda. 

chlamydata Dognin, 1912 (Polyortha); Het- 
eroceres Nouveaux L Amerique du Sud 
6: 50. Holotype ¢, Colombia, Cali, San 
Antonio. 

chloromonas Razowski, 1984 (Chlorortha); 
Acta Zool. Cracov. 27: 215. Holotype 
6, Venezuela, Aragua, Rancho Grande, 
1,100 m, 16—23 October 1966, S. & W. 
Duckworth. 

chortodes Diakonoff, 1968 (Eudemis); Bull. 
U.S. Natl. Mus. 257: 52. Holotype ¢, 
Philippine Islands, Mountain Province, 
Luzon, Baguio, 2 June 1945, J. Francle- 
mont. 

chrysea Heinrich, 1926 (Bactra verutana); 
Bull. U.S. Natl. Mus. 132: 85. Holotype 
6, USA, California, San Bernardino 
Co., Loma Linda, ‘“‘March 24-30.” 

cibdela Razowski, 1988 (Argyrotaenia); 
Acta Zool. Cracov. 31: 409. Holotype 
3d, Colombia, Cusco, Tambomachay, 3 
February 1959, J. Clarke. 

cibriani Miller, 1988 (Rhyacionia); J. Lep- 
id. Soc. 42: 236. Holotype ¢, Mexico, 


VOLUME 102, NUMBER 4 


Mexico, Paso de Cortes, r.f. Pinus har- 
twegii, 12 March 1984, D. Cibrian. 
cinderella Riley, 1872 (Acleris); Board of 
Agric., 4th Annu. Rept. Noxious, Ben- 
eficial and Other Insects . . . of Missou- 
ri: 46. “Type’”” 2, USA, Missouri [no 
data on specimen label], “*928.P.”’ The 
description does not indicate how many 
specimens Riley examined; this is the 
only Riley specimen of A. cinderella we 
could find. Although there is no indi- 
cation in Riley’s hand that is was among 
the original type series, two subsequent 
labels identify it as “‘type.”’ 

cinereolineana Heinrich, 1923 (Thiodia); 
Bull. U.S. Natl. Mus. 123: 52. Holotype 
6, USA, Utah, Juab Co., Eureka, 21 
April 1910, T. Spalding. 

cirrholepida Clarke, 1976 (Icelita); Insects 
of Micronesia 9: 43. Holotype 3, Mi- 
cronesia, Palau, Koror Island, Koror, 25 
April 1957, C. Sabrosky. 

citrana Fernald, 1889 (Tortrix); Entomol. 
Am. 5: 18. ““Type”’ 2, USA, California, 
“the larva had eaten into an orange 
from Coquillett.”’ There is no indication 
of how many specimens Fernald ex- 
amined; this specimen is clearly labeled 
; ype. 

citrogramma Clarke, 1976 (Cryptophlebia); 
Insects of Micronesia 9: 115. Holotype 
?, Micronesia, Kusaie, Hill 541, 165 m, 
18 April 1953, J. Clarke. 

claduncus Razowski, 1988 (Chilips); Acta 
Zool. Cracov. 31: 389. Holotype <4, 
Chile, Centro-Austral, January—March 
1898, V. Izquerdo. 

clarkeana Razowski, 1984 (Polyortha); 
Acta Zool. Cracov. 27: 224. Holotype 
2, Argentina, Tucuman, Ciudad Univ- 
ersitaria, 800 m, 17 February 1959, J. 
Clarke. 

clarkei Blanchard & Knudson, 1983 (Pha- 
neta); Proc. Entomol. Soc. Wash. 85: 
847. Holotype ¢6, USA, Texas, Hem- 
phill Co., Canadian National Grassland, 
Lake Marvin, 9 October 1982, E. Knud- 
son. 


clarkei Diakonoff, 1964 (Bactra); Zool. 


1027 


Verhandel. (Leiden) 70: 8. Holotype 6, 
British Guiana, Vrijheidslust, larva in 
stem of sedge (Cyperus), H. Moore. 

clarkei Obraztsov, 1963 (Acleris); Proc. 
U.S. Natl. Mus. 114: 251. Holotype 6, 
USA, Washington, Kittitas Co., Cle 
Elum, 9 April 1931, J. Clarke. 

clarkei Obraztsov, 1966 (Pseudomeritastis); 
Proc. U.S. Natl. Mus. 118: 226. Holo- 
type 6, Colombia, Cauca, 17 km SE 
Popayan, 2,000 m, 10 January 1959, J. 
Clarke. 

clarki Clarke, 1951 (Corticivora); J. Wash. 
Acad. Sci. 41: 46. Holotype ¢, USA, 
Connecticut, North Guilford, rf. red 
pine (Pinus resinosa), G. Plumb. 

clavana Fernald, 1882 (Semasia); Trans. 
Am. Entomol. Soc. 10: 72. Lectotype 
6, USA, Massachusetts, Truro, August 
8. Designated by Miller (1970). 

clavosa Diakonoff, 1973 (Lobesia); Zool. 
Monogr. Rijksmus. Nat. Hist. 1: 381. 
Holotype 36, Samoa, Fagatogo, Tutuila, 
12 August 1940, ex-Trema(?), O. Swez- 
ey. 

claypoleana Riley, 1882 (Steganoptycha); 
Am. Nat. 16: 914. “Type” @, USA, 
Ohio, em: 2 June 1883. There are two 
identically labeled Riley specimens (i.e., 
‘**360L, Iss. June 2 *83’’), one of which 
has a USNM type label affixed. The 
original description does not indicate 
how many specimens Riley examined. 

clemensiana Fernald, 1879 (Tortrix); Can- 
ad. Entomol. 11: 155. Lectotype d (des- 
ignated here), USA, Maine/Massachu- 
setts/New York/Wisconsin [no locality 
data on specimen label], 10 August 
1880. Selected by Obraztsov in 1956. 
Fernald described this species from 20 
males and 17 females; he did not des- 
ignate a type specimen or a type local- 
ity, instead listing the states from which 
he had specimens. 

clenchi Clarke, 1980 (Proeulia); J. Lepid. 
Soc. 34: 182. Holotype 6, Chile, Des- 
venturadas Islands, San Ambrosio Is- 
land, 450 m, 8 November 1960, G. Kus- 
chel. 


1028 


cneca Obraztsov, 1964 (Proeulia); Proc. 
U.S. Natl. Mus. 116: 193. Holotype ¢, 
Chile, Santiago, Guayacan, 1,100 m, 
October 1952, L. Pefia. 

cockerellana Kearfott, 1907 (Tortrix); 
Trans. Am. Entomol. Soc. 33: 71. Lec- 
totype 6, USA, Colorado, Garfield Co., 
Glenwood Springs, August 1889. Des- 
ignated by Rubinoff and Powell (1999). 

cockleana Kearfott, 1904 (Enarmonia); 
Canad. Entomol. 36: 137. Three syn- 
types as follows: Canada, Alberta, 
Banff, ““VII.9” (2 36) (one labeled 
‘““cotype,”’ the other with a red USNM 
type label); Canada, Manitoba, Awene, 
20 July 1905, Criddle (1 @) (labeled 
““cotype’’). Although Klots (1942) des- 
ignated lectotypes for nearly all other 
Kearfott Olethreutinae in the AMNH, 
he refers to “two paralectotypes”’ of this 
species. 

collilonga Blanchard & Knudson, 1984 
(Pelochrista); Proc. Entomol. Soc. 
Wash. 86: 446. Holotype 6, USA, Tex- 
as, Brown Co., Lake Brownwood State 
Park, 21 April 1966, A. & M. Blan- 
chard. 

coloradanus Fernald, 1882 (Lophoderus); 
Trans. Am. Entomol. Soc. 10: 67. Two 
syntypes (1 6, 1 ¢), USA, Colorado. 
According to the original description, 
this species was described from one 
male and one female from Colorado. 

coloradensis Adamski, 1986 (Apotomis); 
Canad. Entomol. 118: 662. Holotype ¢d, 
USA, Colorado, Maysville, 17 August 
1945, H. Ramstadt. 

coloradensis Heinrich, 1920 (Evetria col- 
jaxiana); Proc. U.S: Natl» Mus: 57: 55. 
Holotype 6, USA, Colorado, Mount 
Manitou, r.f. cones of Abies concolor, 6 
October 1915, J. Pollock, W. Edmon- 
ston, G. Hofer, and A. Champlain. 

comandrana Fernald, 1892 (Teras); Canad. 
Entomol. 24: 121. Lectotype 2, USA, 
Massachusetts, Hampshire Co., Am- 
herst, “‘Hatch Ex. Station.”’ Designated 
by Obraztsov (1963). 

comandranum Clarke, 1953 (Exartema); J. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Wash. Acad. Sci. 43: 230. Holotype 6, 
USA, Illinois, Putnam Co., rf. Coman- 
dra umbellata, 11 June 1942, M. Glenn. 

comstocki Clarke, 1952 (Sonia); Bull. 
South. Calif. Acad. Sci. 51: 62. Holo- 
type ¢d, USA, California, San Diego 
Co., San Felipe Wash, 11 September 
1938, J. Comstock. 

comstockiana Fernald, 1879 (Retinia); Can- 
ad. Entomol. 11: 157. Lectotype’ d, 
USA, New York, Tompkins Co., Ithaca. 
Designated by Miller (1970). 

concitatricana Heinrich, 1923 (Gwendoli- 
na); Bull. U.S. Natl. Mus. 123: 189. Ho- 
lotype 3d, USA, Texas, Kerr Co., Kerr- 
ville, 1 June 1906, E Pratt. 

concubitana Heinrich, 1923 (Gretchena); 
Bull. U.S. Natl. Mus. 123: 181. Holo- 
type 6, USA, Florida, Leon Co., Mon- 
ticello; rf. “Hickoria’ *(\Caryale iS 
March 1914, J. Gill. 

conditana Walsingham, 1879 (Penthina); 
Ill. Lepid. Heter. Brit. Mus. 4: 31. Syn- 
type 6, USA, California, Mendocino 
Co., 24 May 1871. Walsingham de- 
scribed this species from two males; the 
other syntype is presumed to be in the 
BMNH. 

confusa Obraztsov, 1962 (Anopina); Am. 
Mus. Novit. 2082: 30. Holotype 6, 
Mexico, Guerrero, Sierra de las Aguas 
Escondidas, 9,500’, July, H. Smith. 

conigerana Zeller, 1875 (Tortrix); Verhan- 
del. Zool.-Bot. Ges. Wien 25: 227. 
“Type”? 3d, USA, Maine, Packard. Zell- 
er described this species from one male 
from Maine, three females from Mas- 
sachusetts, and one female from New 
York. This is the only specimen of the 
original series that we could find. 

coniogramma Clarke, 1976 (Eucosma); In- 
sects of Micronesia 9: 53. Holotype 4, 
Micronesia, Kusaie, Pukusrik, 9-14 
February 1953, J. Clarke. 

consacculuana Brown, 1991 (Dorithia); in 
Brown and Powell, Univ. Calif. Publ. 
Entomol. 111: 47. Holotype 6, Mexico, 
Chiapas, San Cristébal [de] las Casas, 
17-21 July 1964, P. Spangler. 


VOLUME 102, NUMBER 4 


consobrina Busck, 1914 (Homona); Proc. 
U.S. Natl. Mus. 46: 54. Holotype °&, 
Panama, Porto Bello, March 1911, A. 
Busck. 

consobrinana Heinrich, 1923 (Eucosma); 
Bull. U.S. Natl. Mus. 123: 128. Holo- 
type ¢, USA, South Dakota, Union Co., 
Elk Point, August 1913, C. Ainslie. 

consociana Heinrich, 1923 (Eucosma); 
Bull. U.S. Natl. Mus. 123: 101. Holo- 
type 6, USA, Utah, Juab Co., Eureka, 
23 July 1911, T. Spalding. 

constantia Clarke, 1968 (Cochylis); Proc. 
U.S. Natl. Mus. 125: 22. Holotype 2°, 
Peru, Cusco, Machu Picchu, 2,700 m, 6 
February 1959, J. Clarke. 

constellatana Zeller, 1875 (Sericoris); Ver- 
handel. Zool.-Bot. Ges. Wein 25: 279. 
milype + .d.USA, New «York, 2/71,” 
Speyer. Zeller described this species 
from six males and two females from 
Ohio and New York. This is the only 
Zeller specimen of S. constellatana we 
could find. 

contrasta Brown, 2000 (Lobogenesis); 
Proc. Entomol. Soc. Wash. 102: 32. Ho- 
lotype 6, Bolivia, Incachaca, Cocha- 
bamba, tropical cloud forest area, 2,100 
m, 27 August-5 September 1956, L. 
Pena. 

contrastana Kearfott, 1907 (Commophila); 
Canad. Entomol. 39: 160. Lectotype <, 
USA, Pennsylvania, Allegheny Co., 
Oak Station, 13 June 1907, E Marloff. 
Designated by Klots (1942). 

conversana Walsingham, 1879 (Grapholi- 
tha); Ill. Lepid. Heter. Brit. Mus. 4: 66. 
Syntype 6, USA, Oregon, Camp Wat- 
son, “John Day’s river,” “‘beginning of 
April 1872.’ This species was de- 
scribed from three males and one fe- 
male; the other syntypes presumably are 
in the BMNH. 

coppelia Clarke, 1976 (Icelita tatarana); 
Insects of Micronesia 9: 36. Holotype 
3d, Micronesia, Guam, Piti, 15 Septem- 
ber 1936, rf. Intsia bijuga, O. Swezey. 

cordiae Busck, 1934 (Ancylis); Entomol. 
Am. 13: 154. Holotype 6, Cuba, Ha- 


1029 


vana, Santiago de Vegas, leaf tier on 
Cordia globosa, 26 November 1931, A. 
Otero. 

cornifoliana Riley, 1881 (Phoxopteris); 
Trans. St. Louis Acad. Sci. 4: 324. 
“Type” 3, USA, Kansas, Riley Co., 
Manhattan, r.f. Cornus paniculata, em: 
7 April 1873. Of two specimens re- 
ferred to in the original description, this 
is the only specimen we have found. 

cornucopis Walsingham, 1914 (Enarmon- 
ia); Biol. Centr.-Am., Lepid., Heter. 4: 
240. Holotype 2, Mexico, Oaxaca, Sa- 
lina Cruz, W. Schaus. Described from 
the single female ‘“‘Type.”’ 

cornuta Brown & Obraztsov, 1990 (Cu- 
proxena); in Brown and Powell, Univ. 
Calif. Publ. Entomol. 111: 53. Holotype 
36, Costa Rica, Cartago Province, Juan 
Vinas, W. Schaus. 

cornutana Dyar, 1903 (Epinotia); Proc. En- 
tomol. Soc. Wash. 5: 231. Holotype °, 
USA, Arizona, Coconino Co., Williams, 
“22.7, H. Barber. According to the 
original description, this species was de- 
scribed from a single female. 

cortesi Clarke, 1987 (Cryptophlebia); Acta 
Entomol. Chileana 14: 8. Holotype 6, 
Chile, Iquique Province, Region Tara- 
paca, Valle de Azapa, 25 m, r.f. Acacia 
macracantha, 10 January 1969, R. 
Mendoza. 

corylana Fernald, 1882 (Eccopsis); Trans. 
Am. Entomol. Soc. 10: 71. Lectotype 
3d, USA, New Hampshire, White Moun- 
tains. Designated by Miller (1970). 

corynetes Diakonoff, 1982 (Epinotia); 
Zool. Verhandel. (Leiden) 193: 59. Ho- 
lotype 6, Sri Lanka, Ratnapura District, 
Uggalkaltota, 350’, Irrigation Bunga- 
low, 31 January—8 February 1970, D. 
Davis & B. Rowe. 

cosmocosta Razowski 1987 (Coryssoval- 
va); Tinea 12(suppl.): 130. Holotype ¢, 
Colombia, Cauca, Paramo de Parace, 
Lake San Rafael, 3,570 m, 29 January 
1959; Je Clarke: 

costaricana Razowski 1984 (Histurodes); 
Acta Zool. Cracov. 27: 213. Holotype 


1030 


36, Costa Rica, San Pedro de Montes de 
Oca. 

costimaculana Fernald, 1882 (Penthina); 
Trans. Am. Entomol. Soc. 10: 70. Lec- 
totype 2, USA, Maine, Penobscot Co., 
Orono, 17 June 1881. Designated by 
Miller (1970). 

costinotana Franclemont, 1986 (Lozotaen- 
ia); Proc. Entomol. Soc. Wash. 88: 57. 
Holotype ¢, USA, Maine, Penobscot 
Co., Passadumkeag, 30 June 1956, J. 
Franclemont. 

cracens Diakonoff, 1982 (Eupoecilia); 
Zool. Verhandel. (Leiden) 193: 6. Ho- 
lotype ¢, Sri Lanka, NE District, Kan- 
da-ela Reservoir, 5.6 mi SW Nuwara 
Eliya, 6,200’, 10—21 February 1970, D. 
Davis & B. Rowe. 

crambitana Walsingham, 1879 (Paedisca); 
Ill. Lepid. Heter. Brit. Mus. 4: 43. Syn- 
type 6, USA, California, Mount Shasta, 
August 1871. This species apparently 
was described from one male and one 
female; the female presumably is in the 
BMNH. 

cristata Clarke, 1955 (Orthocomotis); 
Trans. Royal Entomol. Soc. London 
107: 155. Holotype 6, Costa Rica, Ca- 
chi, W. Schaus. 

crocoptila Diakonoff, 1968 (Peridaedala); 
Bull. U.S. Natl. Mus. 257: 79. Holotype 
?, Philippine Islands, Davao Province, 
Mindanao, E slope Mt. McKinley, 
7,000’, 22 September 1946, CNHM 
Philippine Zool. Exped. 1946—47, H. 
Hoogstraal. 

cruentana Blanchard & Knudson, 1981 
(Phaneta); J. Lepid. Soc. 35: 169. Ho- 
lotype 6, USA, Texas, Anderson Co., 
Engeling Wildlife Management Area, 
near Tennessee Colony, 28 June 1978, 
A. & M. Blanchard. 

cryptica Brown, 1991 (Bidorpitia); in 
Brown and Powell, Univ. Calif. Publ. 
Entomol. 111: 75. Holotype 2°, Vene- 
zuela, Aragua, Rancho Grande, 1,100 
m, 1—5 October 1966, S. & W. Duck- 
worth. 

cupressi Heinrich, 1923 (Epinotia hopkin- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


sana); Bull. U.S. Natl. Mus. 123: 207. 
Holotype 6, USA, California, Cypress 
Point, rf. cones of Cupressus macro- 
carpus, 4 November 1915, J. Miller. 

cyanosticha Clarke, 1976 (Cymolomia); In- 
sects of Micronesia 9: 55. Holotype 6, 
Micronesia, Kusaie, Hill 1010, 300 m, 
13 April 1953, J. Clarke. 

cyclopiana Heinrich, 1926 (Polychrosis); 
Bull. U.S. Natl. Mus. 132: 97. Holotype 
2, USA, New Jersey, Burnt Hills, rf. 
seed pods of swamp magnolia, July 
1920, H. Weiss. 

cydna Razowski, 1993 (Apotoforma); Acta 
Zool. Cracov. 36: 185. Holotype 2, 
Venezuela, Rancho Grande, 1,100 m, 
16-23 October 1966, S. & W. Duck- 
worth. 

cylichna Razowski, 1994 (Cylichneulia); 
SHILAP Revta. Lepid. 22: 68. Holotype 
?, Venezuela, Aragua, Rancho Grande, 
15-21 June 1967, 1,100 m, R. Poole. 


daemonicana Heinrich, 1923 (Eucosma); 
Bull. U.S. Natl. Mus. 123: 111. Holo- 
type 6, USA, New Mexico, Manzano 
National Forest, Hell Cafion, “‘flying,”’ 
14 September 1916, C. Heinrich. 

dapsilis Heinrich, 1929 (Eucosma); Proc. 
U.S. Natl. Mus. 75: 5. Holetypecé: 
USA, Wyoming, Park Co., Yellowstone 
National Park, North Park. 

dativa Heinrich, 1928 (Rhyacionia); Proc. 
Entomol. Soc. Wash. 30: 61. Holotype 
36, Japan, Yokahama, r.f. Pinus thunber- 
gii, em: 1 July 1925, A. Kariya. 

davisi Kawabe, 1989 (Archips); Tinea 12: 
191. Holotype 6, Taiwan, Hualien 
Hsien, Tayulin, 2,500 m, 9-18 June 
1980, D. Davis. 

deceptana Busck, 1907 (Pharmacis); J. N. 
Y. Entomol. Soc. 15: 29. Holotype 2, 
USA, Texas, Kerr Co., Kerrville, W. 
Barnes. 

deceptiva Clarke, 1949 (Eulia); Acta Zool. 
Lilloana (Tucuman) 7: 581. Holotype 
?, Brazil, Santa Catarina, Nova Teuton- 
ia, K Plaumann. 

decor Kawabe, 1978 (Enarmonia); Tinea 


VOLUME 102, NUMBER 4 


10: 188. Holotype ¢6, Japan, Honshu, 
Chiba Pref., Tokiwadaira, 5 June 1977, 
A. Kawabe. 

decora Obraztsov, 1966 (Pseudomeritastis); 
Proc. U.S. Natl. Mus. 118: 230. Holo- 
type 6, Bolivia, Cochabamba, Incacha- 
ca, tropical cloud area, 2,100 m, 27 Au- 
gust—5 September 1956, L. Pena. 

decorosa Heinrich, 1929 (Hystricophora); 
Proc. U.S. Natl. Mus. 75: 20. Holotype 
3, USA, Florida, Enterprise, “*4.16.”’ 
Heinrich (1923) consistently misspelled 
Hystrichophora in his revision. 

definitivana Heinrich, 1923 (Anchylopera),; 
Bull. U.S. Natl. Mus. 123: 270. Holo- 
type ¢, USA, Nevada, “July 16—23.”’ 

deflexana Heinrich, 1923 (Epiblema), Bull. 
U.S. Natl. Mus. 123: 144. Holotype ¢, 
USA, Texas, Cameron Co., Browns- 
ville, 27 May 1917, A. Busck. 

delicata Yasuda & Kawabe, 1980 (Croe- 
sia); Tinea 11: 13. Holotype ¢, Japan, 
Nagano Pref., Okutatesina, 23 August 
1963, A. Kawabe. 

delicatana Heinrich, 1923 (Gretchena); 
Bull. U.S. Natl. Mus. 123: 185. Holo- 
type 6, USA, Pennsylvania, Allegheny 
Co., Oak Station, ““May 1-10,” E Mar- 
loff. 

delphinoides Heinrich, 1923 (Thiodia), 
Bull. U.S. Natl. Mus. 123:59. Holotype 
3, USA, Utah, Juab Co., Eureka, 16 
July 1911, T. Spalding. 

delphinus Heinrich, 1923 (Thiodia); Bull. 
U.S. Natl. Mus. 123: 45. Holotype ¢, 
USA, California, Lake Tahoe, Deer 
Park Springs, ‘“‘July 1-7.” 

dendrophila Clarke, 1971 (Dichelopa), 
Smithsonian Contrib. Zool. 56: 105. 
Holotype 6, Rapa Island, Maugaoa, 
950’ (292 m), r.f. Cyathea rapensis, em: 
3 November 1963, J. & T. Clarke. 

deprecatoria Heinrich, 1926 (Olethreutes), 
Bull. U.S. Natl. Mus. 132: 177. Holo- 
type 6, Canada, British Columbia, Wel- 
lington, July, G. Taylor. 

derelicta Heinrich, 1929 (Eucosma); Proc. 
U.S. Natl. Mus. 75: 13. Holotype ¢d, 


1031 


USA, North Carolina, Polk Co., Tryon, 
13 August 1904, Fiske. 

diabolana Blanchard, 1979 (Eucosma),; J. 
Lepid. Soc. 33: 214. Holotype 6, USA, 
Texas, Culberson, Co., Sierra Diablo 
Wildlife Management Area, 6,000’, 31 
March 1970, A. & M. Blanchard. 

diamphidia Clarke, 1968 (Lasiothyris), 
Proc. U.S. Natl. Mus. 125: 49. Holotype 
3, Peru, Cusco, Machu Picchu, 5 Feb- 
ruary 1959, J. Clarke. 

dicaeus Diakonoff, 1968 (Archips); Bull. 
U.S. Natl. Mus. 257: 26. Holotype 6, 
Philippine Islands, Luzon, Mt. Makil- 
ing, Baker. 

digitana Heinrich, 1923 (Epinotia); Bull. 
U.S. Natl. Mus. 123: 215. Holotype 6, 
Canada, British Columbia, Kaslo Creek, 
H. Dyar. 

dilutifuscana Walsingham, 1879 (Sericor- 
is); Ill. Lepid. Heter. Brit. Mus. 4: 33. 
Syntype 6, USA, southern Oregon, 2 
June 1872. Walsingham described this 
species from two males; the other syn- 
type is presumed to be in the BMNH. 

dimorpha Clarke, 1949 (Eulia); Acta Zool. 
Lilloana (Tucuman) 7: 585. Holotype 
?, Brazil, Santa Catarina, 28 March 
1936, E Hoffman. 

dimorphana Barnes & Busck, 1920 (Tor- 
trix); Contrib. Nat. Hist. Lepid. North 
Am. 4: 215. Holotype 6, Canada, Brit- 
ish Columbia, Victoria. 

discobola Diakonoff, 1968 (Strepsicrates),; 
Bull. U.S. Natl. Mus. 257: 85. Holotype 
36, Philippine Islands, Davao Province, 
Mindanao, E slope Mt. McKinley, 
7,200’, 10 September 1946, mossy 
stunted forest, CNHM Philippine Ex- 
ped. 1946-47, H. Hoogstraal. 

discretivana Heinrich, 1921 (Eucosma); J. 
Agric. Res. 20: 823. Holotype ¢, USA, 
Texas, Harris Co., Sheldon, 10 April 
1919, Johnson. 

dispersa Brown, 1990 (Auratonota), Flori- 
da Entomol. 73: 154. Holotype 6, Pan- 
ama, Cocle Province, Valle, 22 April 
1965, S. & W. Duckworth. 

disputabilis Obraztsov 1963 (Acleris); Proc. 


1032 


U.S. Natl. Mus. 114: 262. Holotype <6, 
Canada, British Columbia, Goldstream, 
16 October 1902. 

dissitana Grote, 1879 (Ptycholoma); North 
Am. Entomol. 1: 29. “Type”? 2, USA, 
4 July 1878. This specimen, labeled 
‘“‘type’’ by someone other than Grote, is 
the only Grote specimen of P. dissitana 
we could locate. The original descrip- 
tion does not indicate how many spec- 
imens were examined. 

distincta Obraztsov, 1966 (Pseudomeritas- 
tis); Proc. U.S. Natl. Mus. 118: 228. 
Holotype 2, Costa Rica, Cartago Prov- 
ince, La Florida, 500’, W. Schaus. 

dognini Obraztsov, 1966 (Pseudatteria); 
Proc. U.S. Natl. Mus. 118: 588. Holo- 
type 6d, Ecuador, Environs of Loja, 
1887. 

dominica Brown, 1993 (Auratonota); Pan- 
Pac. Entomol. 69: 314. Holotype 6, 
West Indies, Dominica, 1.7 mi E Point 
Casse, light trap, 24 March 1965, W. 
Wirth. 

domna Clarke, 1968 (Amallectis); Proc. 
U.S. Natl. Mus. 125: 34. Holotype 6, 
Colombia, Narifo, Volcan Galeras, 
3,000 m, 14 January 1959, J. Clarke. 

donaldana Kawabe, 1993 (Notocelia); Tin- 
ea 13: 238. Holotype 6, Taiwan, Nan- 
tou Hsien, Mei-feng, 30 km S Tayuling, 
2,200 m, 1—8 July 1980, D. Davis. 

doria Clarke, 1976 (Laspeyresia); Insects 
of Micronesia 9: 123. Holotype 6, Mi- 
cronesia, Palau, Babelthuap, Ngarhe- 
long, 1 May 1957, C. Sabrosky. 

dorsalana Dyar, 1903 (Tortrix); Proc. En- 
tomol. Soc. Wash. 5: 231. Syntypes (4 
3636, 2 22), USA, Arizona, Coconino 
Co., Williams, rf. oak, em: 14 June 
1901, A. Schwartz. According to the 
original description, Dyar described this 
species from four males and four fe- 
males; all of the syntypes cited above 
are labeled “Type No. 6736 USNM,”’ 
consistent with the original description. 

dorsata Clarke, 1988 (Dichelopa); Smith- 
sonian Contrib. Zool. 416: 133. Holo- 
type 2, Marquesas Islands, Hiva Oa, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Mt. Feani, 3,400’, 1 March 1968, J. & 
T. Clarke. 

druana Walsingham, 1914 (Tortrix); Biol. 
Centr.-Am., Lepid., Heter., 4: 288. Ho- 
lotype ¢, Mexico, Durango, r.f. cynipid 
gall on Quercus, em: 11 February 1897. 
This species was described from the sin- 
gle male. 

dryocremna Meyrick, 1932 (Polyortha); 
Exotic Microlepidoptera 4: 343. Lecto- 
type ¢ (designated here), Guatemala, 
Palin, Schaus & Barnes. Selected by J. 
Clarke. Meyrick’s description indicates 
*“6 ex. [examples] (type U.S. Nat. 
Muss): 

duckworthorum Brown, 1991 (Cuproxena); 
in Brown and Powell, Univ. Calif. Publ. 
Entomol. 111: 67. Holotype 3, Costa 
Rica, Cartago Province, Turrialba, 17— 
21 February 1965, S. & W. Duckworth. 

dulciana Heinrich, 1923 (Gretchena); Bull. 
U.S. Natl. Mus 123: 182. Holotype 6, 
USA, New Jersey, Greenwood Lake, 
“*V.30,”” W. Kearfott. 

dysmorphia Clarke, 1968 (Lasiothyris); 
Proc. U.S. Natl. Mus. 125: 47. Holotype 
3, Bolivia, Cochabamba, Incachaca, 
tropical cloud area, 2,100 m, 27 Au- 
gust—5 September 1956, L. Pefia. 


eburata Heinrich, 1929 (Eucosma); Proc. 
U.S. Natl. Mus. 75: 6. Holotype ¢, 
USA, Arizona, Mohave Co., “Aug. 24— 
Sey 

edemoidana Dyar, 1903 (Eucosma); Proc. 
Entomol. Soc. Wash. 5: 229. Two syn- 
types (2 22), USA, Arizona, Coconino 
Co., Williams, ‘*19-7,’’ W. Barnes; 
USA, New Mexico, Las Vegas HS, 
18.8." According to the original de- 
scription, this species was described 
from the two females cited above. 

egens Razowski, 1999 (Netechma); Polskie 
Pismo Entomol. 68: 97. Holotype 6, 
Colombia, Narifo, Volcan Galeras, 
2,900 m, 13 January 1959, J. Clarke. 
Razowski (1999) incorrect identifies the 
disposition of the type as NHMW. 

elaborata Kawabe, 1976 (Pseudohedya); 


VOLUME 102, NUMBER 4 


Tinea 10: 45. Holotype 6, Japan, Ao- 
mori Pref., Tuta Spa, 20 July 1957, A. 
Kawabe. 

eleonora Obraztsov, 1962 (Anopina); Am. 
Mus. Novit. 2082: 12. Holotype ¢, 
USA, Arizona, Apache Co., Alpine, 6 
June 1937, G. & J. Sperry. 

elitha Clarke, 1976 (Heleanna physalodes); 
Insects of Micronesia 9: 19. Holotype 
36, Micronesia, Ponape Island, Colonia, 
13 January 1953, J. Clarke. 

elongana Brown, 1990 (Cuproxena); in 
Brown and Powell, Univ. Calif. Publ. 
Entomol. 111: 54. Holotype ¢, Vene- 
zuela, Amazonas (territory), Cerro de 
Neblina Camp VII, 1,850 m, 0°5S1’N, 
65°58'W, 2—4 December 1984, Brown. 

embaphion Razowski, 1984 (Saphenista),; 
Ann. Zool. Warsaw 38: 276. Holotype 
6, Venezuela, Mérida, 8 km SE Apar- 
taderos, 22 February 1976, C. & O. 
Flint. 

embolina Razowski, 1984 (Saphenista); 
Ann. Zool. Warsaw 38: 277. Holotype 
2, Venezuela, Mérida, 4 km S Santo 
Domingo, 19-23 February 1976, C. & 
O. Flint. 

emera Razowski, 1993 (Acleris); Acta 
Zool. Cracov. 36: 191. Holotype 6, Bo- 
livia, Cochabamba, Incachaca, 27 Au- 
gust—5 September 1956, L. Pena. 

emigratella Busck, 1909 (Amorbia), Proc. 
Entomol. Soc. Wash. 11: 201. Syntype 
6, Hawaiian Islands, Oahu, Tantulus, 
O. Swezey. Busck did not indicate how 
many specimens he examined. 

enucleata Razowski, 1999 (Netechma); 
Polskie Pismo Entomol. 68: 95. Holo- 
type 6, Colombia, Narifio, Volcan Ga- 
leras, 300 m, 14 January 1959, J. 
Clarke. Razowski (1999) incorrect iden- 
tifies the disposition of the type as 
NHMW. 

episticta Clarke, 1949 (Eulia); Acta Zool. 
Lilloana (Tucuman) 7: 584. Holotype 
6, Brazil, Santa Catarina, Nova Teuton- 
ia, September 1948, E Plaumann. 

equadora Brown, 1990 (Paraptila); J. Lep- 


1033 


id. Soc. 44: 271. Holotype 6, Ecuador, 
Shell-Mera, 18 April 1958, R. Hodges. 

erigeronana Riley, 1881 (Conchylis),; 
Trans. St. Louis Acad. Sci. 4: 316. Ho- 
lotype 6, USA, Texas, Columbia, rf. 
‘‘cecidomyidous gall on Erigeron,”’ em: 
24 February 1879, E. Schwarz. Riley 
described this species from a single 
specimen. 

erotella Heinrich, 1923 (Carpocapsa), 
Proc. Entomol. Soc. Wash. 25: 121. Ho- 
lotype 2, USA, Maryland, Montgom- 
ery Co., Hyattsville, rf. Pinus taeda, 
em: 22 May 1915, A. Busck. 

erubesca Kawabe, 1978 (Gypsonoma); Tin- 
ea 10: 187. Holotype 6, Japan, Hok- 
kaido, Sapporo, Matuyama Park, 30 
July 1950, A. Mutsuura. 

escharia Clarke, 1976 (Trymalitis); Insects 
of Micronesia 9: 142. Holotype 3d, Mi- 
cronesia, Guam, Ritidian, 2 August 
1945, J. Gressitt. 

essigana Busck, 1929 (Amorbia); Calif. 
Dept. Agric. Mon. Bull. 18: 276. Ho- 
lotype 6, USA, California, San Diego 
Co., Chula Vista, rf. avocado leaves, 8 
December 1928, R. McLean. 

ethnica Heinrich, 1923 (Epinotia); Bull. 
U.S. Natl. Mus. 123: 201. Holotype ¢, 
USA, California, San Diego Co., San 
Diego, “‘June 16—23.” 

euchaldera Clarke, 1955 (Orthocomotis); 
Trans. Royal Entomol. Soc. London 
107: 146. Holotype ¢, Colombia, June 
1909, Fassel. 

eupista Diakonoff, 1968 (Stenarchella); 
Bull. U.S. Natl. Mus. 257: 99. Holotype 
3, Philippine Islands, Davao Province, 
Mindanao, La Lum Mt., Calian, 5,000’, 
29-21 December 1920, Clagg, CM 
Acc. 9163. 

evestigana Razowski, 1984 (Polyortha), 
Acta Zool. Cracov. 27: 221. Holotype 
6, El Salvador, Cerro Miramundo, 
2,300 m, 11 January 1969, S. Steinhau- 
ser. 

exacerbatricana Heinrich, 1923 (Epible- 
ma); Bull. U.S. Natl. Mus 123: 146. Ho- 


1034 


lotype 6, USA, North Carolina, South- 
ern Pines, ““Aug. 16—23.” 

exaeresimum Heinrich, 1926 (Exartema); 
Bull. U.S. Natl. Mus. 132: 160. Holo- 
type 6, USA, Texas, Dallas Co., Dallas. 

excerptionana Heinrich, 1923 (Eucosma); 
Bull. U.S. Natl. Mus. 123: 117. Holo- 
type 6, USA, Nevada, Washoe Co., 
Verdi, ‘“‘June 1 to 10,’’ A. Vachell. 

exclusoriana Heinrich, 1923 (Eucosma); 
Bull. U.S. Natl. Mus. 123: 110. Holo- 
type d, USA, Texas, La Salle Co., Co- 
tulla, 12 May 1906, Crawford & Pratt. 

excusabilis Heinrich, 1923 (Eucosma); 
Bull. U.S. Natl. Mus. 123: 123. Holo- 
type 6, USA, California, Lake Tahoe, 
Deer Park Springs. 

exolivata Clarke, 1955 (Orthocomotis); 
Trans. Entomol. Soc. London 107: 148. 
Holotype @, Brazil, Santa Catarina, 
Nova Teutonia, August 1936, H. Parish. 

exomilana Franclemont, 1986 (Lozotaenia); 
Proc. Entomol. Soc. Wash. 88: 58. Ho- 
lotype ¢, USA, North Carolina, Macon 
Co., Highlands, 3,865’, 11 July 1958, J. 
Franclemont. 

exoristus Razowski, 1988 (Exoletuncus); 
Acta Zool. Cracov. 31: 390. Holotype 
6, Colombia, Cauca, Paramo de Parce, 
Lake San Rafael, 3,570 m, 29 January 
1959, J. Clarke. 

expolitana Heinrich, 1923 (Eucosma); Bull. 
U.S. Natl. Mus. 123: 132. Holotype 6, 
USA, Utah, Utah Co., Provo, 11 August 
1908, T. Spalding. 

exulis Issiki & Stringer, 1932 (Gnorismo- 
neura),; Stylops 1: 125. Holotype 6, 
Taiwan, Taihoku, 18 October 1923, S. 
Issiki. 


famula Zeller, 1875 (Teras tristana); Ver- 
handel. Zool.-Bot. Ges. Wien 25: 214. 
Lectotype 6, North America [no col- 
lecting data]. Designated by Obraztsov 
(1963). Zeller’s original description re- 
fers to a male, consistent with the spec- 
imen cited above. However, the associ- 
ated Busck genitalia slide (Dec 6 1924; 
USNM 25944) is of a female, and Ob- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


raztsov’s (1963: 236) lectotype desig- 
nation also refers to the specimen as fe- 
male. 

fernaldana Grote, 1880 (Paedisca); N. Am. 
Entomol.: 98. “‘Type’” 6, USA, Colo- 
rado. This is the only Grote specimen 
of fernaldana we could locate; it bears 
a labeling identifying it as the type, but 
not in Grote’s hand. The original de- 
scription does not indicate how many 
specimens were examined. 

fernaldana Walsingham 1879 (Cochylis); 
Ill. Lepid. Heter. Brit. Mus. 4: 27. Syn- 
type 6, USA, northern California, Sep- 
tember 1871. This species was de- 
scribed from four males and one female; 
all of the other syntypes are presumed 
to be in the BMNH. 

ferreana Busck, 1915 (Sparganothis); Proc. 
Entomol. Soc. Wash. 17: 86. Holotype 
2, USA, New York, Herkimer Co., II- 
ion, 10 July 1912, H. McElhose. 

ferruginana Fernald, 1882 (Semasia); 
Trans. Am. Entomol. Soc. 10: 72. Lec- 
totype 2, USA, Massachusetts, Good- 
ell. Designated by Miller (1970). 

ferrugineana Riley, 1881 (Exartema); 
Trans. St. Louis Acad. Sci.. 4: 317. 
“Type’’ 3, USA, Missouri, St. Louis 
Co., St. Louis, rf. leaves of plum. Riley 
described this species from two males, 
only one of which could be located by 
us. 

ferruginiguttana Fernald, 1882 (Teras); 
Trans. Am. Entomol. Soc. 10: 65. 
“Type”? 2, USA, Colorado. Of two fe- 
males that comprised the type series, we 
could find only one. It is represented by 
a portion of the thorax bearing a single 
leg and a small piece of the left fore- 
wing. 

ferruginus Pogue, 1986 (Apolychrosis); in 
Cibridn-Tovar et al., Cone and Seed In- 
sects of the Mexican Conifers: 21. Ho- 
lotype ¢, Mexico, Tlaxcala, Municipio 
de Terrenate, Villareal, rf. Pseudotsuga 
macrolepis, em: 18 August—6 Septem- 
ber 1981, J. Mendez & R. Campos. 

fertoriana Heinrich, 1923 (Thiodia); Bull. 


VOLUME 102, NUMBER 4 


U.S. Natl. Mus. 123: 264. Holotype 6, 
Canada, British Colombia, Goldstream, 
**10-5-03.” 

filiana Busck, 1907 (Hendecaneura); J. N. 
Y. Entomol. Soc. 15: 135. Holotype 6, 
USA, California, Riverside Co., West 
Riverside, 26 October 1905. 

finitimana Heinrich, 1923 (Kundrya); Bull. 
U.S. Natl. Mus. 123: 192. Holotype 6, 
USA, New Hampshire, Rockingham 
Co., Hampton, 11 June 1909, S. Shaw. 

fishiana Fernald, 1882 (Peronea); Trans. 
Am. Entomol. Soc. 10: 66. Lectotype 
6, USA, Maine, Penobscot Co., Orono, 
17 September 1879. Designated by Ob- 
raztsov (1963). 

flavana Fernald, 1905 (Eucosma pergan- 
deana); Canad. Entomol. 37: 399. Lec- 
totype 6, USA, Texas. Designated by 
Miller (1970). 

flavibasana Fernald, 1882 (Cenopis); Trans. 
Am. Entomol. Soc. 10: 69. Lectotype 3 
(designated here), USA, Illinois. Select- 
ed by Powell in 1965. Although there is 
little doubt that the lectotype is one of 
2 specimens referred to in the original 
description, Fernald misidentified the 
sex, indicating two females, one from 
Texas and one from Illinois. 

flavifasciana Kawabe, 1976 (Olethreutes),; 
Tinea 10: 47. Holotype 2, Japan, Gun- 
ma Pref., Kuridaira, 20 July 1951, M. 
Hoshino. 

flavillana Dyar, 1903 (Epinotia); Proc. En- 
tomol. Soc. Wash. 5: 230. Three syn- 
types (1 6, 2 22), USA, Arizona, Co- 
conino Co., Williams, bred from flower 
stalk, June. According to the original 
description, this species was described 
from three specimens. 

flexicostalis Dognin, 1908 (Tortrix); Ann. 
Soc. Entomol. Belgium 52: 32. Holo- 
type ¢, Peru (SE), Oconeque, Cara- 
baya, ‘10.07,’ Warren. 

flexura Clarke, 1986 (Dichelopa); Smith- 
sonian Contrib. Zool. 416: 124. Holo- 
type 6, Marquesas Islands, Hiva Oa, 
Feani, 3,800’, 20 February 1968, J. & 
T. Clarke. 


1035 


flintana Brown, 1991 (Cuproxena); in 
Brown and Powell, Univ. Calif. Publ. 
Entomol. 111: 69. Holotype 2, Mexico, 
Jalisco, Rt. 94, km 42, N Chapala [4 mi 
toward Guadalajara from Chapala; top 
of hill], 16-18 July 1966, O. Flint & A. 
Ortiz. 

footiana Fernald, 1882 (Eccopsis); Bull. 
Buffalo Soc. Nat. Sci. 4: 53. Lectotype 
2, USA, New York. Designated by 
Miller (1970). 

formosanus Kawabe, 1987 (Archippus); 
Tinea 7: 122. Holotype ¢, Taiwan, Chi- 
ayi, Alishan, 2,200 m, 9-11 July 1964, 
H. Inoue. 

foxcana Kearfott, 1907 (Phalonia); Trans. 
Am. Entomol. Soc. 33: 84. Lectotype 
36, USA, Ohio, Hamilton Co., Cincin- 
nati, 18 August 1903, A. Braun. Kear- 
fott described this species from three 
specimens from Cincinnati, Ohio, and 
Plummers Island, Maryland, with no in- 
dication of which locality was the 
source of two of the three. Klots (1942) 
designated the above-mentioned lecto- 
type, indicating that there were two lec- 
toparatypes in the AMNH: a female 
from Plummers Island and another 
specimen without an abdomen (he 
didn’t provide its collection data). Al- 
though he correctly cited the date of 
collection of the lectotype, he incorrect- 
ly cited its collection locality as Plum- 
mers Island. 

fragariae Walsh & Riley, 1869 (Anchylo- 
pera); Am. Entomol. 1: 89. Lectotype 
2, USA, Illinois, Whiteside or Bureau 
Co. According to the original descrip- 
tion, this species was described from 
nine specimens, only two of which 
could be located by us. Designated by 
Miller (1973). 

fragariana Busck, 1919 (Tortricodes); 
Proc. Entomol. Soc. Wash. 21: 52. Ho- 
lotype 6, Canada, British Columbia, 
Victoria, 17 August 1918. 

frangula Clarke, 1968 (Amallectis); Proc. 
U.S. Natl. Mus. 125: 30. Holotype °, 
Venezuela, Aragua, Rancho Grande, 


1036 


1,100 m, 16—23 October 1966, S. & W. 
Duckworth. 

fraternana Busck 1907 (Hendecaneura); J. 
N. Y. Entomol. Soc. 15: 134. Holotype 
6, USA, California, Riverside Co., 
West Riverside, October. 

fratruelis Heinrich, 1923 (Eucosma); Bull. 
U.S. Natl. Mus. 123: 98. Holotype <4, 
USA, North Carolina, Southern Pines, 
8-15 August. 

fraudabilis Heinrich, 1923 (Eucosma); 
Bull. U.S. Natl. Mus. 123: 98. Holotype 
3, USA, North Carolina, Southern 
Pines. According to the original descrip- 
tion, the type series was collected be- 
tween 1 June and 23 July. 

fritillana Blanchard & Knudson, 1981 (Eu- 
cosma); J. Lepid. Soc. 35: 170. Holo- 
type 6, USA, Texas, Anderson Co., En- 
geling Wildlife Management Area, near 
Tennessee Colony, 28 June 1978, A. 
Blanchard. 

frustrana W. Comstock, 1880 (Retinia); 
Rept. U.S. Dept. Agric. 1879: 236. Lec- 
totype 2, USA, Virginia, near Washing- 
ton (D.C.), “Tortrix on P. inops,”’ em: 
18 June 1879. Designated by Miller 
(1967). 

fucana Walsingham, 1879 (Lozotaenia); Il. 
Lepid. Heter. Brit. Mus. 4: 12. Syntype 
3, USA, Oregon (southern), May 1872. 
This species was described from four 
males; the other syntypes are presum- 
ably in the BMNH. 

fullerea Riley, 1869 (Penthina); J. Horti- 
cult., Boston, Oct. 1868: 12. ““Type”’ 2, 
USA, “seed pods of Antherrinum.”” An 
associated label indicates that the spec- 
imen is an unpublished lectotype se- 
lected by Miller in 1982. Riley did not 
indicate how many specimens he ex- 
amined. 

fulvomixtana Kawabe, 1974 (Zeiraphera); 
Tyo to Ga 25: 98. Holotype d, Japan, 
Gunma Pref., Doaiguchi, 12 July 1972, 
S. Shimeki. 

fumipennis Dognin, 1904 (Atteria); Ann. 
Soc. Entomol. Belgium 48: 133. Holo- 
type 6, Colombia, Micay, Aoiat, 1869. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


fumosana Powell, 1978 (Rhyacionia); in 
Powell and Miller, U.S. Dept. Agric., 
Agric. Handb. 514: 21. Holotype 4, 
USA, Colorado, El Paso Co., Colorado 
Springs, Rock Creek Canyon, 2 May 
1959, M. May. 

fumoviridana Heinrich, 1923 (Epinotia); 
Bull. U.S. Natl. Mus. 123: 208. Holo- 
type 3d, USA, California, Siskiyou Co., 
Shasta Retreat, “Aug. 16-23.” 

fuscalbana Zeller, 1876 (Sericoris); Verhan- 
del. Zool.-Bot. Ges. Wein 25: 284. 
“Type”? 6, USA, [Maine or Massachu- 
setts according to the original descrip- 
tion], Packard. This is the only Zeller 
specimen of fuscalbana that we were 
able to locate. The original description 
does not indicate how many specimens 
were examined, but it does indicate that 
the series included only males. 

fuscana Barnes & Busck, 1920 (Peronea); 
Contrib. Nat. Hist. Lepid. North Am. 4: 
216. Holotype ¢6, Canada, Manitoba, 
Aweme, 18 April 1905, Criddle. 

fuscodorsana Kearfott, 1904 (Commophi- 
la); Canad. Entomol. 36: 141. Holotype 
36, Canada, British Colombia, Kaslo, 7 
June 1903, J. Cockle. Klots (1942) cor- 
rectly stated that the holotype is in the 
USNM, consistent with the original de- 
scription. 

fuscomaculatus Brown, 2000 (Odonthali- 
tus); Proc. Entomol. Soc. Wash. 102: 
42. Holotype 2, Mexico, Michoacan, 
San Lorenzo, Rt. 15, km 206, 19 July 
1966, O. Flint & A. Ortiz. 


galena Clarke, 1971 (Tritopterna); Smith- 
sonian Contrib. Zool. 56: 124. Holotype 
3, Rapa Island, Teumukopuke, 500’, 7 
October 1963, J. & T. Clarke. 

galerasiana Razowski, 1988 (Uncicida); 
Acta Zool. Cracov. 31: 396. Holotype 
3, Colombia, Narifio, Volcan Galeras, 
2,900 m, 13 January 1959, J. Clarke. 

gallaesaliciana Riley, 1881 (Grapholitha); 
Trans. St. Louis Acad. Sci. 4: 320. 
“Type” 2, USA, Missouri, St. Louis 
Co., St. Louis, ‘“‘Lepidopterous gall on 


VOLUME 102, NUMBER 4 


willow,” 17 June 1873. This is one of 
two specimens cited by Riley in the 
original description; the location of the 
other is unknown. 

gambra Razowski, 1988 (Ernocornutina); 
Acta Zool. Cracov. 31: 399. Holotype 
36, Argentina, Tucuman, Ciudad Univ- 
ersitaria, 800 m, 17 February 1959, J. 
Clarke. 

gampsognathos Razowski, 1988 (Gaurun- 
cus); Acta Zool. Cracov. 31: 405. Ho- 
lotype 6, Bolivia, Cochabamba, Inca- 
chaca, tropical cloud area, 2,100 m, 27 
August—5 September 1956, L. Pefia. 

garai Miller, 1987 (Gretchena); J. Lepid. 
Soc. 41: 151. Holotype 2, Ecuador, 4 
km S Loja, rf. Juglans neotropica, 28 
June 1986, A. Samaniego. 

gatesclarkei Kawabe, 1992 (Acleris); Tinea 
13: 171. Holotype 6, Taiwan, Nantou 
Hsien, Tsuifeng, 2,400 m, 29 December 
1989, A. Kawabe. 

gelastes Razowski, 1988 (Gauruncus); Acta 
Zool. Cracov. 31: 404. Holotype d, Ar- 
gentina, Tucuman, Ciudad Universitar- 
ia, 800 m, 17 February 1959, J. Clarke. 

gemellana Heinrich, 1923 (Eucosma cir- 
culana); Bull. U.S. Natl. Mus. 123: 96. 
Holotype 6, USA, Florida, Hillsbor- 
ough Co., St. Petersburg, May. 

gentilii Brown, 1998 (Argentulia); J. Lepid. 
Soc. 52: 180. Holotype ¢, Argentina, 
Neuquén, Paso Cordoba, 1,300 m, 20 
February 1980, M. Gentili. 

georgiella Hulst, 1887 (Myelois); Entomol. 
Am. 3: 136. “Type” 6, USA, Colora- 
do. This is the only Hulst specimen we 
could find of M. georgiella; it is labeled 
““type.”’ The original description does 
not indicate how many specimens were 
examined. 

gerda Busck, 1911 (Olethreutes); Proc. 
U.S. Natl. Mus. 40: 227. Holotype 2, 
French Guiana, St. Jean, Maroni, W. 
Schaus. 

giganteana Riley, 1881 (Paedisca); Trans. 
St. Louis Acad. Sci. 4: 318. “‘Pseudo- 
type” ¢, USA, Missouri, “‘Barlon.” 
According to Riley’s original descrip- 


1037 


tion, this species was described from 2 
females, one from ‘‘Kansas (G. E Gau- 
mer)’ and one from “Iowa, March, 
1874 (Hoffmeister).’’ Although not in- 
cluded in the original series, the speci- 
men from Missouri bears a type label in 
Riley’s hand and represents a potential 
neotype if one is deemed necessary. 

gigantica Busck, 1920 (Hysterosia); Insect. 
Inscit. Menst. 8: 87. Holotype 2, Mex- 
ico, Distrito Federal, Mexico City, R. 
Muller. 

gilletteana Dyar, 1903 (Eucosma); Proc. 
Entomol. Soc. Wash. 5: 229. Syntype 6, 
USA, Arizona, Coconino Co., Williams, 
16.6; foun syntypes. (Gi Gc. 2), 
USA, Colorado. According to Dyar, this 
species was described from the series (n 
= 5) cited above; all of the specimens 
bear red labels “Type No: 6737 
U.S.N.M.” 

gloriola Heinrich, 1931 (Eucosma); Proc. 
Entomol. Soc. Wash. 33: 196. Holotype 
3, USA, Connecticut, Stamford, B.T.R. 
Lab colony, rf. white pine tip, 6 May 
193i" 

gnoma Clarke, 1986 (Dichelopa); Smith- 
sonian Contrib. Zool. 416: 138. Holo- 
type 6, Marquesas Islands, Fatu Hiva, 
Hanavave, 10’, 12 April 1968, J. & T. 
Clarke. 

goodelliana Fernald, 1882 (Phoxopteris); 
Trans. Am. Entomol. Soc. 10: 69. Lec- 
totype 6, USA Maine, New Hampshire, 
or Massachusetts [no locality data on 
specimen labels]. Designated by Miller 
(1970). 

graceana Powell, 1960 (Argyrotaenia); 
Pan-Pac. Entomol. 36: 93. Holotype <6, 
USA, California, San Bernardino Co., 
San Bernardino Mts., Hathaway Creek, 
2 August 1940, C. Henne. 

grandicula Heinrich, 1926 (Laspeyresia); 
Bull. U.S. Natl. Mus. 132: 54. Holotype 
3, USA, Virginia, Giles Co., Mountain 
Lake, 14—21 June 1907. A. Braun. 

grandiflavana Walsingham, 1879 (Paedis- 
ca); Ill. Lepid. Heter. Brit. Mus. 4: 50. 
Syntype ¢, USA, California, Lake Co., 


1038 


17-19 June 1871. This species was de- 
scribed from one male and two females; 
the other syntypes presumably are in the 
BMNH. 

grandis Busck, 1907 (Phalonia); J. N. Y. 
Entomol. Soc. 15: 23. Holotype 2°, 
USA, Colorado, Jefferson Co., Golden, 
Chimney Gulch, Oslar, 1 July 1904. 

gratiana Kawabe, 1974 (Hedya); Tyo to Ga 
25: 101. Holotype ¢, Japan, Honmura, 
Kuchinoerabujima Island, 14 August 
1973, r.f. Glochidon obovatum, A. Ka- 
wabe. 

gratuitana Heinrich, 1923 (Epiblema); 
Bull. U.S. Natl. Mus. 123: 268. Holo- 
type 6, USA, Washington, East Sound, 
11 July 1901. 

graziella Blanchard, 1968 (Eucosma); J. 
Lepid. Soc. 22: 143. Holotype ¢, USA, 
Texas, Brewster Co., Big Bend National 
Park, Green Gulch, 11 October 1966, A. 
& M. Blanchard. 

grindeliana Busck, 1906 (Cydia); Canad. 
Entomol. 38: 211. Two syntypes (1 6, 
1 2), USA, Texas, Clarendon, r.f. Grin- 
delia squarrosa, 28 September 1905, 
em: 4 October 1905, Hunter. In the orig- 
inal description, Busck identified the 
type as labeled “‘Type #9804”’; howev- 
er, both of the specimens listed above 
have identical red type labels with this 
number. 

griselda Blanchard & Knudson, 1981 (Eu- 
cosma); J. Lepid. Soc. 35: 173. Holo- 
type 6, USA, Texas, Brewster Co., Big 
Bend National Park, Chisos Basin, 7 
April 1967, A. & M. Blanchard. 

groteana Fernald, 1882 (Cenopis); Trans. 
Am. Entomol. Soc. 10: 69. Holotype 2, 
USA, Ohio. This species was described 
from a single specimen. 

guerrerana Obraztsov, 1964 (Anopina); 
Am. Mus. Novit. 2082: 15. Holotype ¢, 
Mexico, Guerrero, Sierra de las Aguas 
Escondidas, 9,500’, July, H. Smith. 

guiana Busck, 1913 (Olethreutes); Insect. 
Inscit. Menst. 1: 92. Holotype 6, British 
Guiana [Guyana], Georgetown, “larvae 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


feed on a common weed in the cane 
field.” 

gunniana Busck, 1907 (Phalonia); J. N. Y. 
Entomol. Soc. 15: 26. Holotype 2°, 
USA, Maryland, Montgomery Co., 
Plummers Island, September 1903, A. 
Busck. 

guttulana Blanchard, 1979 (Eucosma); J. 
Lepid. Soc. 33: 214. Holotype ¢, USA, 
Texas, Kennedy Co., Padre Island Na- 
tional Seashore, 19 July 1976, A. & M. 
Blanchard. 

gyraleus Diakonoff, 1982 (Archips); Zool. 
Verhandel. (Leiden) 193: 87. Holotype 
6, Sri Lanka, Ratnapura District, Ug- 
galkaltota, 350’, Irrigation Bungalow, 
31 January—8 February 1970, Davis & 
Rowe. 


habrosana Heinrich, 1923 (Exentera); Bull. 
U.S. Natl. Mus. 123: 178. Holotype 6, 
USA, California, San Diego Co., San 
Diego, 17 March 1912, W. Wright. 

hadrotes Clarke, 1986 (Dichelopa); Smith- 
sonian Contrib. Zool. 416: 126. Holo- 
type 6, Marquesas Islands, Nuku Hiva, 
Tunoa Ridge, 2,900’, 23 January 1968, 
J. & T. Clarke. 

hasseanthi Clarke, 1952 (Eucosma); Bull. 
South. Calif. Acad. Sci. 52: 60. Holo- 
type 6, USA, California, Orange Co, 
‘“‘bred from Hasseanthus variegatus,” 
23 August 1936, T. Hower. 

heindelana Fernald, 1905 (Acleris); Am. 
Nat. 39: 870. Lectotype ¢, Canada, 
Manitoba, Winnipeg, A. W. Hanham. 
Designated by Obraztsov (1963). 

heliocausta Dognin, 1912 (Atteria); Heter- 
oceres Nouveaux de L Amerique du Sud 
6: 51. Holotype 6, Colombia, Cali, San 
Antonio, 2,000 m, 22 September 1908, 
Fassel. 

helianthana Riley, 1881 (Grapholitha); 
Trans. St.Louis’ Acad’ Scite=t-ws13: 
“Type” 6, USA, Texas, “gall on sun- 
flower,” 12 August 1873. Riley de- 
scribed this species from two speci- 
mens, only one of which has been lo- 
cated by us. 


VOLUME 102, NUMBER 4 


hemeropis Dognin, 1912 (Olethreutes); 
Heteroceres Nov. L Amerique du Sud 6: 
49. Holotype 6, Colombia, Cali, San 
Antonio, Fassel. 

hemitephras Clarke, 1976 (Lasiothyris); 
Proc. U.S. Natl. Mus. 125: 51. Holotype 
2, Mexico, Puebla, Tehuacan, 10 Oc- 
tober. 

henicodes Razowski, 1988 (Bicavernaria); 
Acta Zool. Cracov. 31: 400. Holotype 
36, Peru, Cusco, Machu Picchu, 2,385 
m, 6 February 1959, J. Clarke. 

hennei Clarke, 1947 (Eucosma); Bull. 
South. Calif. Acad. Sci. 46: 51. Holo- 
type 6, USA, California, Los Angeles 
Co., El Segundo sand dunes, em: 3 Oc- 
tober 1940, C. Henne. 

heos Razowski, 1988 (Helicteulia); Acta 
Zool. Cracov. 31: 388. Holotype 6, Bo- 
livia, Cochabamba, Incachaca, tropical 
cloud area, 2,100 m, 27 August—5 Sep- 
tember 1956; 1: "Pena: 

herbacea Clarke, 1955 (Orthocomotis); 
Trans. Royal Entomol. Soc. London 
107: 151. Holotype 6, Costa Rica, San 
Pedro de Montes de Oca, r.f. avocado, 
em: 15 January 1933, C. Ballou. 

herbaria Busck, 1920 (Sociphora); Insect. 
Inscit. Menst. 8: 85. Holotype 6, Gua- 
temala, Cuyuga, W. Schaus. 

heterophaea Clarke, 1968 (Phalonidia); 
Proc. U.S. Natl. Mus. 125: 36. Holotype 
6, Colombia, Antioquia, La Estrella, 
1,730 m, 13 December 1959, FE Luis 
Gallego M. 

heucherana Heinrich, 1923 (Epinotia); 
Bull. U.S. Natl. Mus. 123: 217. Holo- 
type 6, USA, Virginia, Arlington Co., 
Rosslyn, larvae mining leaves of ‘“‘alum 
root’? [Heuchera americana], C. Hein- 


rich. 
hieroglyphana Blanchard & Knudson, 1984 
(Grapholita); Proc. Entomol. Soc. 


Wash. 86: 448. Holotype ¢, USA, Tex- 
as, Culberson Co., Guadalupe Moun- 
tains, Nickel Creek, 10 July 1968, A. & 
M. Blanchard. 

hieroglypta Walsingham, 1914 (Olethreu- 
tes); Biol. Centr.-Am. Lepid., Heter. 4: 


1039 


250. Holotype 6, Mexico, Veracruz, 
Orizaba, W. Schaus. Walsingham clear- 
ly designated a “‘Type”’ male deposited 
in the USNM. 

hipeana Grote, 1876 (Conchylis); Canad. 
Entomol. 8: 207. Holotype ¢, Canada, 
Ontario, Port Stanley, W. Saunders. Al- 
though the specimen bears a label 
‘“‘London, Ont.,”” according to the orig- 
inal description it was taken at Port 
Stanley, by ““W. Saunders from London, 
Ontario.”” This species was described 
from a single male. 

hiranoi Kawabe, 1980 (Gypsonoma); Tinea 
11: 27. Holotype 6, Japan, Nagano 
Pref., Nakakaruizawa, 14 June 1964, A. 
Kawabe. 

hiroshii Kawabe, 1980 (Zeiraphera); Tinea 
11: 26. Holotype 6, Japan, Gunma 
Pref., Kumaneta’ra, 14 July 1952, M. 
Hoshino. 

hirsutana Walsingham, 1879 (Paedisca); 
Ill. Lepid. Heter. Brit. Mus. 4: 50. Syn- 
type 6, USA, California, Sonoma Co., 
23 May 1871. This species was de- 
scribed from four males; the other three 
presumably are in the BMNH. 

hodgesi Heppner, 1989 (Argyrotaenia); 
Florida Entomol. 72: 102. Holotype ¢, 
USA, Florida, Glade Co., Fisheating 
Creek, 7-10 May 1964, R. Hodges. 

hodgesi Razowski, 1993 (Apotoforma); 
Acta Zool. Cracov. 36: 186. Holotype 
?, Panama, Cerro Campana, 11—14 July 
i967, O>S. Fling 

hodsoni Miller, 1986 (Pseudexentera); J. 
Lepid. Soc. 40: 223. Holotype 3, USA, 
Pennsylvania, Allegheny Co., Oak Sta- 
tion, 10 April 1910, E Marloff. 

hoffmanana Brown, 1991 (Cuproxena); in 
Brown and Powell, Univ. Calif. Publ. 
Entomol. 111: 60. Holotype 2, Brazil, 
Santa Catarina, Neu-Bremen, 12 June 
1931, E Hoffman. 

hohuanshana Kawabe, 1986 (Zeiraphera); 
Entomol. Pap. Pres. Kurosawa, Tokyo: 
79. Holotype ¢, Taiwan, Hualien Hsien, 
Hohuanshan, 3,100 m, 30 July—1 Au- 
gust 1983, A. Kwabe. 


1040 


hohuanshana Kawabe, 1989 (Acleris); Tin- 
ea 12: 194. Holotype 3, Taiwan, Hu- 
alien Hsien, Hohuanshan, 3,100 m, 30 
July—1 August 1983, A. Kawabe. 

hohuanshanensis Kawabe, 1985 (Clepsis); 
Tinea 12: 5. Holotype ¢, Taiwan, Hu- 
alien Hsien, Hohuanshan, 3,100 m, 30 
July—1 August 1983, A. Kawabe. 

holographa Clarke, 1965 (Nesochoris); 
Proc. U.S. Natl. Mus. 117: 74. Holotype 
6, Chile, Masatierra, Plazoleta del 
Yunque; 200%m=s 9). February “1952; P. 
Kuschel. 

homolopa Diakonoff, 1968 (Eudemis); 
Bull. U.S. Natl. Mus. 257: 54. Holotype 
2, Philippine Islands, Davao Province, 
Minandao, 5717. 

horii Kawabe, 1987 (Cryptophlebia); Tinea 
12: 141. Holotype 6, Japan, Okinawa 
Island, Okubi, Kin, rf. Bruguiera gym- 
norrhyza, 8 May 1983, S. Hori. 

hoshinoi Kawabe, 1964 (Hastula); Tyo to 
Ga 15: 2. Holotype 6, Japan, Honshu, 
Tokyo, Setagaya, 16 June 1963, A. Ka- 
wabe. 

hostilis Diakonoff, 1956 (Bactra); Zool. 
Verhandel. (Leiden) 29: 57. Holotype 
3d, Japan, Honshiu, Funakoshi, Yoko- 
suka, 11 September 1953. 

houseri Miller, 1959 (Petrova); Ohio J. Sci. 
59: 230. Holotype 6, USA, Ohio, 
Washington Co., Veto, r.f., Pinus echin- 
ata, 18 June 1954, W. Miller. 

huachucensis Obraztsov, 1961 (Argyro- 
taenia montezumae); Am. Mus. Novit. 
2048: 7. Holotype ¢, USA, Arizona, 
Cochise Co., Huachuca Mtns., August 
1905, H. Skinner. 

hubbardana Busck, 1907 (Phalonia); J. N. 
Y. Entomol. Soc. 15: 27. Holotype 2, 
USA, Arizona, Pima Co., Tucson, rf. 
Koeberlinia spinosa, em: 12 May 1897, 
H. Hubbard. 

hypericana Ely, 1910 (Peronea); Proc. En- 
tomol. Soc. Wash. 12: 68. Holotype 2, 
USA, Virginia, Fairfax Co., Great Falls, 
r.f., Hypericum prolificum, em: 17 June 
1909, C. Ely. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


icelitodes Diakonoff, 1982 (Eucosma); 
Zool. Verhandel. (Leiden) 193: 45. Ho- 
lotype 6, Sri Lanka, Mannar District, 
Olaithoduvai, 10 mi NW of Mannar, 0— 
50’, 4—5 November 1976, G. Hevel, R. 
Dietz, S. Karunaratne & D. Balasooriya. 

icogramma Clarke, 1968 ([razona); Proc. 
U.S. Natl. Mus. 125: 38. Holotype 2°, 
Guatemala, Volcan Santa Maria, Octo- 
ber, Schaus and Barnes. 

illotana Walsingham, 1879 (Paedisca); Ill. 
Lepid. Heter. Brit. Mus. 4: 48. Two syn- 
types (1 6, 1 2), Oregon, Rogue River, 
May 1872. Walsingham described this 
species from two males and one female; 
the other male syntype is presumed to 
be in the BMNH. 

imbrica Kawabe, 1978 (Hedya); Tinea 10: 
181. Holotype ¢, Japan, Yakushima Is- 
land, Shiratani, 14 September 1972, T. 
Watanabe. 

imbridana Fernald, 1905 (Cydia); Canad. 
Entomol. 37: 400. Lectotype 6, USA, 
Kansas, Pottawatomie Co., Onaga, F 
Crevecoeur. Designated by Miller 
(1970). 

imitativa Heinrich, 1926 (Grapholitha); 
Bull. U.S. Natl. Mus. 132: 34. Holotype 
36, USA, California, San Francisco. 

implicata Heinrich, 1924 (Proteoteras); J. 
Wash. Acad. Sci. 14: 390. Holotype 6, 
USA, Florida, Dade Co., Everglades, 
ex-larva, “April 16—23.” 

implicata Heinrich, 1931 (Thiodia); Proc. 
U.S. Natl. Mus. 79: 7. Holotype ¢, 
USA, Washington, Rochester, 13 June 
1929, W. Baker. 

improvisana Heinrich, 1923 (Epinotia); 
Bull. U.S. Natl. Mus. 123: 269. Holo- 
type 6, USA, California, Tulare Co., 
Mineral King, “‘June 16—23.” 

incognita Obraztsov, 1963 (Acleris); Proc. 
U.S. Natl. Mus. 114: 254. Holotype °, 
USA, Idaho, Moscow Mountains, 7 Au- 
gust 1933, J. Clarke. 

inconspicua Obraztsov, 1964 (Proeulia); 
Proc. U.S. Natl. Mus. 116: 190. Holo- 
type 6, Chile, Santiago, La Obra, Oc- 
tober 19525 L.jRena: 


VOLUME 102, NUMBER 4 


indagatricana Heinrich, 1923 (Thiodia); 
Bull. U.S. Natl. Mus. 123: 56. Holotype 
3, USA, Utah, Utah Co., Provo, 26 Au- 
gust 1908, T. Spalding. 

indentanus Dyar, 1903 (Phthinolophus); 
Proc. Entomol. Soc. Wash. 5: 306. 
Twenty-nine syntypes from various lo- 
calities in the eastern United States. A 
female from USA, Virginia, Fortress 
Munroe, 24 June 1884, H. Dyar, was 
placed in the type collection, presum- 
ably by J. Clarke; the rest of the syn- 
types are in the main collection. Ac- 
cording to the original description, this 
species was described from 17 males 
and 21 females. 

indigena Yasuda, 1978 (Eurydoxa); Trans. 
Lepid. Soc. Jap. 29: 119. Holotype <6, 
Taiwan, Rengwati, 29 March 1929, S. 
Issiki. 

infelix Heinrich, 1923 (Epiblema); Bull. 
U.S. Natl. Mus. 123: 151. Holotype 6, 
USA, North Carolina, Polk Co., Tryon, 
25 May 1904, Fiske. 

infernalis Heinrich, 1920 (Commophila); 
Proc. U.S. Natl. Mus. 57: 61. Holotype 
3, USA, New Mexico, Manzano Natl. 
Forest, Hell Canyon, rf. berries of Jun- 
iperus, 31 June 1917, C. Heinrich. 

infida Heinrich, 1926 (Aphania); Bull. U.S. 
Natl. Mus. 132: 121. Holotype 3, Can- 
ada, Quebec, St. Johns Co., St. Therese 
Island, 9 June 1915, W. Chagnon. 

infimbriana Dyar, 1904 (Thiodia); Proc. 
U.S. Natl. Mus. 27: 927. Holotype °, 
Canada, British Columbia, Kaslo, H. 
Dyar. 

influana Heinrich, 1923 (Thiodia); Bull. 
U.S. Natl. Mus. 123: 49. Holotype ¢, 
USA, California, Siskiyou Co., Shasta 
Retreat, “July 1-7.” 

infuscata Heinrich, 1923 (Endothenia); 
Proc. Entomol. Soc. Wash. 25: 109. Ho- 
lotype ¢6, USA, Maryland, Forest Glen, 
10 July 1914, O. Heidemann. 

ingens Heinrich, 1926 (Laspeyresia); Bull. 
WS: NatliMus: 132:63:) Holotype ¢; 
USA, Florida, Hillsborough Co., St. Pe- 
tersburg. 


1041 


ingrata Heinrich, 1926 (Laspeyresia); Bull. 
U.S. Natl. Mus. 132: 50. Holotype ¢, 
Canada, Manitoba, Aweme, 25 May 
1905, Criddle. 

injectiva Heinrich, 1926 (Hedulia); Bull. 
U.S. Natl. Mus. 132: 65. Holotype ¢d, 
USA, Nevada, Reno, rf. cones of Pinus, 
em: 24 January 1911, J. Smith. 

inopinata Heinrich, 1928 (Laspeyresia); 
Proc. Entomol. Soc. Wash. 30: 91. Ho- 
lotype ¢, China, South Manchuria, Kin- 
shu, on apple, 4 August 1927, T. Kondo. 

inopiosa Heinrich, 1926 (Laspeyresia); 
Bull. U.S. Natl. Mus. 132: 46. Holotype 
2, USA, Idaho, Kootenai Co., Coeur 
d’Alene, rf. Pinus contorta, em: 11 
May 1916, J. Evendon. 

inoueit Kawabe, 1968 (Pandemis); Tinea 7: 
121. Holotype 36, Taiwan, Chiayi Coun- 
ty, Alishan, 2,200 m, 9-11 July 1964, 
H. Inoue. 

inouei Kawabe, 1972 (Eupoecilia); Tinea 9: 
250. Holotype 6, Japan, Kagosaka- 
toge, base of Mt. Fuji, 13 August 1969, 
H. Inoue. 

inouei Kawabe, 1987 (Metendothenia); Tin- 
ea 12: 139. Holotype ¢, Japan, Mie 
Pref., Hokusei-machi, Otsujishinden, 8 
May 1986, T. Mano. 

insidiosana Heinrich, 1923 (Epiblema); 
Bull. U.S. Natl. Mus. 123: 145. Holo- 
type 6, USA, North Carolina, Southern 
Pines, “‘June 1-7.” 

insignata Heinrich, 1924 (Thiodia); J. 
Wash. Acad. Sci. 14: 386. Holotype ¢, 
USA, Colorado, San Juan Co., Silver- 
ton, “July 8-15” [genitalia slide only; 
pinned specimen has not been found]. 

insignis Heinrich, 1928 (Petrova); Proc. 
Entomol. Soc. Wash. 30: 63. Holotype 
3, Japan, Yokohama, rf. Pinus thun- 
bergii, em: 27 August 1924. 

inspersa Heinrich, 1931 (Phaecasiophora); 
Proc. U.S. Natl. Mus. 79: 13. Holotype 
3, USA, Florida, Hillsborough Co., St. 
Petersburg, 3 March 1915, R. Ludwig. 

insulanus Kawabe, 1965 (Archippus); Tyo 
to Ga 16: 23. Holotype 3, Japan, Oki- 


1042 
noerabu-jima Is., 8 April 1957, M. 
Umebayashi. 

interruptolineana Fernald, 1882 (Penthina); 
Trans. Am. Entomol. Soc. 10: 70. Lec- 
totype 6, USA, New Hampshire or 
Massachusetts [no locality data on spec- 
imen labels]. Designated by Miller 
(1970). 

invidana Barnes & Busck, 1920 (Tortrix); 
Contrib. Nat. Hist. Lepid. North Am. 4: 
215. Holotype 6, Canada, British Co- 
lumbia, Vancouver Island, Duncans, 
Hanham. 

iodes Clarke, 1968 (Hysterosia); Proc. U.S. 
Natl. Mus. 125: 4. Holotype 6, Guate- 
mala, Volcan Santa Maria, Schaus & 
Barnes. 

iresinephora Razowski, 1988 (Eriotortrix); 
Acta Zool. Cracov. 31: 403. Holotype 
3d, Colombia, Bogota, Chico, 23 Janu- 
ary 1959, J. Clarke. 

iridana Barnes & Busck, 1920 (Platynota); 
Contrib. Nat. Hist. Lepid. North Am. 4: 
212. Holotype 2, USA, Florida, Hills- 
borough Co., St. Petersburg, June. 

irroratana Walsingham, 1879 (Paedisca); 
Ill. Lepid. Heter. Brit. Mus. 4: 48. Syn- 
type 6, USA, California, Mendocino 
Co., 7-10 June 1871. Walsingham de- 
scribed this species from five males; all 
but the USNM syntype are presumed to 
be in the BMNH. 

isipida Razowski, 1988 (Eriotortrix); Acta 
Zool. Cracov. 31: 403. Holotype 3, Co- 
lombia, Bogota, Chico, 23 January 
1959, J. Clarke. 

issikii Kawabe, 1980 (Aterpia); Tinea 11: 
17. Holotype 3, Japan Honmura, Ku- 
chinoarabu Island, 27—31 July 1973, A. 
Kawabe. 

issikii Razowski, 1977 (Hysterosia); Tyo to 
Ga 28: 35. Holotype 2, Japan, Kuway- 
ama, 23 June 1916. 

ivana Fernald, 1901 (Tortrix); J. N. Y. En- 
tomol. Soc. 9: 51. Holotype 6, USA, 
Florida, rf. Iva imbricata, 10 March 
1900. This species was described from 
a single male. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


jJaponensis Kawabe, 1980 (Cydia); Tinea 
11: 28. Holotype ¢, Japan, Akita Pref., 
Onuma Spa, Hachimantai, 9-10 August 
1974, R. Sato. 

Japonica Kawabe, 1978 (Eucoenogenes); 
Tinea 10: 185. Holotype 3, Japan, Hon- 
shu, Gunma Pref., Mt. Mikaboyama, 2 
August 1968, S. Shimeki. 

Jenningsi Powell, 1978 (Rhyacionia); in 
Powell and Miller, U.S. Dept. Agric., 
Agric. Handb. 514: 17. Holotype 6, 
USA, Arizona, Sitgreaves National For- 
est, r.f. Pinus ponderosa, D. Jennings. 

Jinboi Kawabe, 1965 (Clepsis); Kontyu 34: 
459. Holotype 6, Japan, Honshu, South 
Alps, Sanpuku-goya, | August 1964, K. 
Jinbo [‘‘Jimbo”’ on label]. 

Jinboi Kawabe, 1976 (Cymolomia); Tinea 
10: 44. Holotype ¢, Japan, Hokkaido, 
Mt. Daisetsu (Kurodake), 16 August 
1969, K. Jinbo. 

jJordani Clarke, 1955 (Orthocomotis); 
Trans. Royal Entomol. Soc. London 
107: 142. Holotype ¢, Brazil, New Bre- 
men, 11 August 1936, EK Hoffman. 

juglandana Fernald, 1879 (Tortrix); Canad. 
Entomol. 11: 155. Two syntypes (1 6, 
1 2), USA, Massachusetts/Ohio/Wis- 
consin, and Canada (Ontario) [no local- 
ity data on specimen labels]. Fernald 
described this species from 11 males 
and 15 females; he did not designate a 
type specimen or a type locality, instead 
listing the states from which he had 
specimens. 

juncticiliana Walsingham, 1879 (Paedisca); 
Ill. Lepid. Heter. Brit. Mus. 4: 75. Two 
syntypes (2 6¢), USA, California, 
Shasta Co., 16 July 1871. This species 
was described from seven males; the 
other five presumably are in the BMNH. 


kana Busck, 1906 (Hemimene); Proc. Biol. 
Soc. Wash. 19: 182. Lectotype ¢, Can- 
ada, British Columbia, Kaslo, H. Dyar. 
Designated by Miller (1983). 

khasiensis Miller, 1977 (Petrova); J. Lepid. 
Soc. 31: 135. Holotype 6, India, Assam 


VOLUME 102, NUMBER 4 


(Meghalaya), Upper Shillong, March 
1963. 

kincaidiana Fernald, 1900 (Phoxopteris); in 
Dyar, Proc. Wash. Acad. Sci. 2: 500. 
Holotype 6, USA, Alaska, Metlakahtla, 
“6.4.99,” Harriman Expedition ‘99, T. 
Kincaid. 

kirishimensis Kawabe, 1974 (Eudemopsis); 
Kontyu 42: 390. Holotype ¢, Japan, 
Kagoshima Pref., Mt. Kirishima [Kari- 
sima], 30 June 1958, A. Kawabe. 

kiyosatoensis Kawabe, 1980 (Endothenia); 
Tinea 11: 20. Holotype ¢, Japan, Ya- 
mamshi Pref., Kiyosato, 1,300 m, 25— 
28 August 1971, A. Kawabe. 

knudsoni Miller, 1986 (Pseudexentera); J. 
Lepid. Soc. 40: 224. Holotype 6, USA, 
Texas, Kleberg Co., Riviera Beach site, 
24 February 1984, E. Knudson. 

koebelei Obraztsov, 1959 (Aphelia); Am. 
Mus. Novit. 1964: 7. Holotype 6, USA, 
Washington, Easton, Koebele. 


komonana Kearfott, 1907 (Hysterosia); 


Canad. Entomol. 37: 121. Lectotype 6, 
USA, California, Santa Clara Co. Des- 
ignated by Klots (1942). 

kurosawai Kawabe, 1986 (Notocelia); En- 
tomol. Pap. Pres. Kurosawa, Tokyo: 80. 
Holotype 6, Taiwan, Hualien Hsien, 
Hohuanshan, 3,100 m, 30 July—1 Au- 
gust 1983, A. Kawabe. 

kusaiensis Clarke, 1976 (Dudua aprobola); 
Insects of Micronesia 9: 86. Holotype 
6, Micronesia, Kusaie, Mutunlik, 14 
February 1953, J. Clarke. 

kuscheli Clarke, 1980 (Proeulia); J. Lepid. 
Soc. 34: 184. Holotype ¢, Chile, Des- 
venturadas Islands, San Ambrosia Is- 
land, 450 m, on flowers of Thamnoseris 
lacerata, 14 November 1960, G. Kus- 
chel. 

kusunokii Kawabe, 1993 (Apotomis); Tyo to 
Ga 43: 257. Holotype 6, Japan, Keun- 
shita, Mt. Daisetsu, 28 July 1983. 


laciniana Zeller, 1875 (Phoxopteris); Ver- 
handel. Zool.-Bot. Ges. Wein 25: 253. 
“Type’’ 6, USA, Massachusetts, 1871. 
Of the three males and one female re- 


1043 


ferred to in the original description, this 
is the only specimen we have been able 
to locate. 

laetitia Clarke, 1968 (Cochylis); Proc. U.S. 
Natl. Mus. 125: 25. Holotype ¢, Argen- 
tina, Tucuman, Ciudad Universitaria, 
800 m, 20 February 1959, J. Clarke. 

lagopana Walsingham, 1879 (Steganopty- 
cha); Ill. Lepid. Heter. Brit. Mus. 4: 71. 
Syntype 2, USA, California, Colusa 
Co., 28 June 1871. This species was de- 
scribed from two males and three fe- 
males; presumably all but the above 
mentioned syntype are in BMNH. 

lamberti Franclemont, 1986 (Sparganoth- 
is); Proc. Entomol. Soc. Wash. 88: 56. 
Holotype 3d, USA, South Carolina, 
Oconee Co., Cherry Hill Recreation 
Area, Route 107, 2,000’, 22 August 
1958, J. Franclemont. 

lambertiana Busck, 1915 (Tortrix); Proc. 
Entomol. Soc. Wash. 17: 86. Holotype 
3, USA, Oregon, Jackson Co., Ashland, 
rf. Pinus lambertii, em: 27 June 1914, 
P. Sergent. Although the original de- 
scription gives “‘Oakland, Oreg.”’ as the 
type locality, the label on the specimen 
indicates Ashland. 

lantana Busck, 1910 (Crocidosema); Proc. 
Entomol. Soc. Wash. 12: 132. Holotype 
36, Hawaii, Oahu, Tantalus, O. Swezey. 

larana Brown, 2000 (Lobogenesis); Proc. 
Entomol. Soc. Wash. 102: 29. Holotype 
3d, Venezuela, Lara, Yacumba National 
Park, 13 mk SE Sanare, 4,800’, cloud 
forest, blacklight, 4-7 March 1978, J. 
Heppner. 

largo Heppner, 1981 (Cydia); J. Lepid. Soc. 
35: 278. Holotype 36, USA, Florida, 
Monroe Co., Key Largo, 15 mi NE Key 
Largo City, 16 June 1974, J. Heppner. 

laricana Busck, 1916 (Laspeyresia); Proc. 
Entomol. Soc. Wash. 18: 152. Holotype 
2, USA, Montana, Evaro, r.f., Larix oc- 
cidentalis, May 1914, J. Brunner. 

lariciana Kawabe, 1980 (Zeiraphera); Tin- 
ea 11: 24. Holotype 6, Japan, Nagano 
Pref., Shig-k6gen, 28 June 1964, Oza- 
wa. 


1044 


latens Heinrich, 1929 (Thiodia); Proc. U.S. 
Natl. Mus. 75: 2. Holotype 6, USA, 
California, Tulare Co., Monachee 
Meadows, 8,000’, “July 8-14.” 

lathami Forbes, 1937 (Eucosma); J. N. Y. 
Entomol. Soc. 43: 131. Holotype <4, 
USA, New York, Long Island, Orient, 
18 June 1935, R. Latham. 

latiana Brown, 1991 (Cuproxena); in 
Brown and Powell, Univ. Calif. Publ. 
Entomol. 111: 52. Holotype ¢, Vene- 
zuela, Aragua, Rancho Grande, 1,100 
m, 10 May 1967, L. Rodriguez & C. 
Andara. 

laticurva Heinrich, 1929 (Eucosma); Proc. 
U.S. Natl. Mus. 75: 4. Holotype ¢, 
USA, California, Sierra Nevada. 

latipunctana Walsingham, 1879 (Cochylis); 
Ill. Lepid. Heter. Brit. Mus. 4: 29. Syn- 
type 6, USA, California, Mendocino 
Co., 31 May 1871. This species was de- 
scribed from two males; the other syn- 
type presumably is in the BMNH. 

lautana Powell, 1960 (Argyrotaenia); Pan- 
Pac. Entomol. 36: 90. Holotype <6, 
USA, California, San Bernardino Co., 
San Bernardino Mtns., Camp Baldy, 
“June 24-30.” 

lavana Busck, 1907 (Phalonia); J. N. Y. 
Entomol. Soc. 15: 27. Holotype ¢, 
USA, Maryland, Montgomery Co., Hy- 
attsville, A. Busck. 

leguminana Busck, 1907 (Phalonia); J. N. 
Y. Entomol. Soc. 15: 28. Holotype ¢, 
USA, Washington, D.C., rf. Gleditchia 
horrida, Feburary. 

leguminis Heinrich, 1943 (Laspeyresia); 
Proc. Entomol. Soc. Wash. 45: 71. Ho- 
lotype 6, Peru, rf. beans, 19 August 
1930, Willie. 

leopardana Busck, 1906 (Hemimene); Proc. 
Biol. Soc. Wash. 19: 181. Holotype 2, 
USA, Maryland, Montgomery Co., Hy- 
attsville, June 1906, A. Busck. 

lepida Heinrich, 1924 (Epinotia cruciana); 
J. Wash. Acad. Sci. 14: 391. Holotype 
36, USA, New Hampshire, Coos Co., 
Mt. Washington, 4,000’, ‘“‘July 24-31.” 

leucobasis Busck, 1916 (Laspeyresia); 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Proc. Entomol. Soc. Wash. 18: 152. Ho- 
lotype 6, USA, Montana, Evaro, r.f. La- 
rix occidentalis, May 1914, J. Brunner. 

leucognoma Clarke, 1976 (Eumarissa); In- 
sects of Micronesia 9: 32. Holotype 6, 
Micronesia, Guam, Mt. Alifan, ex-um- 
bellifer(?), 21 May 1936, O. Swezey. 

leucothorax Clarke, 1955 (Orthocomotis); 
Trans. Royal Entomol. Soc. London 
107: 150. Holotype 6, Brazil, New Bre- 
men, 6 June 1936, E Hoffman. 

lindana Fernald, 1892 (Steganoptycha); 
Canad. Entomol. 24: 178. Lectotype d, 
Canada, Ontario, Hamilton. Designated 
by Miller (1970). 

lindseyana Obraztsov, 1962 (Choristoneura 
lambertiana); Am. Mus. Novit. 2101: 
16. Holotype 6, USA, California, Mo- 
doc Co., Warner Mtns., 3 mi E Davis 
Creek, 15-23 July 1922, 5,500’, A. 
Lindsey. 

lineana Fernald, 1901 (Eucosma); J. N. Y. 
Entomol. Soc. 9: 50. Holotype 2, USA, 
Florida, Palm Beach Co., Palm Beach, 
r.f. Anona laurifolia, H. Dyar. This spe- 
cies was described from a single female. 

linitipunctana Blanchard & Knudson, 1983 
(Phaneta); J. Lepid. Soc. 37: 140. Ho- 
lotype 6, USA, Texas, Nueces Co., 
North Padre Island, 9 September 1974, 
A. & M. Blanchard. 

listerana Kearfott, 1907 (Pharmacia); 
Trans. Am. Entomol. Soc. 33: 80. Lec- 
totype 6 (designated here), USA, Penn- 
sylvania, Lackawanna Co., Nicholson, 6 
July 1904, A. Lister. Selected by Obraz- 
tsov in 1953. 

lobata Razowski, 1988 (Argyrotaenia); 
Acta Zool. Cracov. 31: 408. Holotype 
3d, Bolivia, Cochabamba, Incachaca, 
tropical cloud area, 2,100 m, 27 Au- 
gust-5 September 1956, L. Pena. 

loricana Grote, 1880 (Phoxopteris); Canad. 
Entomol, 12: 218. ‘‘Type*> 62sAe 
Ohio, Montgomery Co., Dayton, G. Pi- 
late. According to the original descrip- 
tion, the “‘type is in the collection of 
Fernald,’’ which almost certainly refers 
to the specimen cited above. 


VOLUME 102, NUMBER 4 


louisiana Busck, 1907 (Phalonia); J. N. Y. 
Entomol. Soc. 15: 24. Holotype 6, 
USA, Missouri, St. Louis Co., 2 mi W 
St. Louis, August 1904, A. Busck. 

luciferana Kawabe, 1980 (Zeiraphera); 
Tinea 11: 26. Holotype 6, Japan, Mi- 
yagi Pref., Sakunmi, 22 June 1969, T. 
Watanabe. 

luctuosana Blanchard, 1979 (Epiblema); J. 
Lepid. Soc. 33: 184. Holotype ¢, USA, 
Texas, Nueces Co., N. Padre Island, 6 
April 1978, A. & M. Blanchard. 

luculentana Heinrich, 1920 (Evetria); Proc. 
U.S. Natl. Mus. 57: 56. Holotype ¢, 
USA, Colorado, El Paso Co., r.f. Pinus 
scopulorum, 5 May 1916, W. Edmon- 
ston. 

luoyingensis Kawabe, 1992 (Acleris); Tinea 
13: 175. Holotype 6, Taiwan, Hualien 
Hsien, Houhuanshan, Luoying Lodge, 
2,800 m, 31 December 1988, A. Ka- 
wabe. 

lupicinia Clarke, 1971 (Dichelopa); Smith- 
sonian Contrib. Zool. 56: 113. Holotype 
36, Rapa Island, Pt. Maraia, rf. Rumex 
crispus, em: 26 October 1963, J. & T. 
Clarke. 


macdunnoughi Obraztsov, 1963 (Acleris); 
Proc. U.S. Natl. Mus. 114: 214. Holo- 
type 2, USA, Massachusetts, Worcester 
Co., Winchendon, 29 September 1902. 

machimiana Barnes & Busck, 1920 (Spar- 
ganothis); Contrib. Nat. Hist. Lepid. 
North Am. 4: 211. Holotype °, USA, 
Arizona, Cochise Co., Paradise, July. 

macswaini Powell, 1980 (Decodes); Pacific 
Insects 22: 89. Holotype 3, Mexico, 
Nuevo Leon, 3 mi E Galeana, 5,000’, 
7-9 August 1963, W. Duckworth & D. 
Davis. 

maculana Fernald, 1901 (Lipoptycha); J. N. 
Y. Entomol. Soc. 9: 51. Holotype ¢, 
USA, Florida, Palm Beach Co., Palm 
Beach, r.f. Schoepfia arborescens, em: 
24 February 1900, H. Dyar. There are 
three specimens in the USNM collec- 
tion with virtually identical collecting 
data, two of which bear labels in Fer- 


1045 


nald’s hand indicating “‘type’’; the latter 
two are undoubtedly the two males that 
comprise the original series. Although 
Miller (1970) was unable to find the ho- 
lotype, one of the males bears a red 
USNM type label (“‘Type 5413”’’) con- 
sistent with the original description, im- 
plicating it as the holotype. 

maculatana Walsingham, 1879 (Paedisca); 
Ill. Lepid. Heter. Brit. Mus. 4: 48. Two 
syntypes (2) 22), USA, California; 
Lake Co., 17-19 June 1871. This spe- 
cies was described from three males and 
three females; the other syntypes pre- 
sumably are in the BMNH. 

maenamii Kawabe, 1974 (Olethreutes); Tyo 
to Ga 25: 102. Holotype ¢, Japan, Izu 
Island, Shikinejima, 16 June 1966, T. 
Maenami. 

magnifica Razowski & Becker, 1999 (Au- 
ratonota); Revta. Bras. Zool. 16: 1174. 
Holotype 6, Venezuela, Aragua, Ran- 
cho Grande, 1,100 m, at light in mon- 
tane tropical forest, 16 June 1973, J. C. 
& K. G. Schaffer. 

magnoliana Fernald, 1892 (Cacoecia); 
Canad. Entomol. 24: 121. ‘“‘Type” 6, 
USA, New York, Tompkins Co., Ithaca, 
C.U. Exp. No. 292, rf. Magnolia acu- 
minata, 22 June 1891, M. Slingerland. 
Fernald did not indicate how many 
specimens he examined. 

maiana Kearfott, 1907 (Phalonia); Trans. 
Am. Entomol. Soc. 33: 82. Lectotype 
36, USA, New Jersey, Essex Co., Essex 
County Park, 9-14 May 1910, W. Kear- 
fott. Designated by Klots (1942). 

maiorina Heinrich, 1923 (Bactra); Proc. 
Entomol. Soc. Wash. 25: 105. Holotype 
3, USA, Virginia, Arlington Co., Ar- 
lington, rf. Scirpus fluviatilis, 1 July 
1920. 

malana Fernald, 1882 (Eccopsis); Trans. 
Am. Entomol. Soc. 10: 72. Lectotype 
36, USA, New York, Kings Co., Brook- 
lyn, J. Smith. Designated by Miller 
(1970). 

manilkara Heppner, 1981 (Dichrorampha); 
Florida Entomol. 64: 274. Holotype 6, 


1046 


USA, Florida, Monroe Co., Middle 
Torch Key, 12 June 1974, r.f. Manilkara 
emarginata, em: 6 July 1974, J. He- 
ppner. 

manoi Kawabe, 1987 (Hedya); Tinea 12: 
140. Holotype 6, Japan, Mie Pref., 
Hokushei-machi, Otsujishenden, 8 May 
1986, T. Mano. 

mareda Clarke, 1955 (Orthocomotis); 
Trans. Royal Entomol. Soc. London 
107: 142. Holotype 6, Brazil, Nova 
Teutonia, May 1938, E Plaumann. 

mariana Fernald, 1882 (Lophoderus); 
Trans. Am. Entomol. Soc. 10: 67. Lec- 
totype 6 (designated here), USA, 
Maine, Penobscot Co., Orono. Selected 
by Obraztsov 1956. According to the 
original description, Fernald had four 
males from Maine (Orono), Massachu- 
setts, and New York. 

maritima Dyar, 1904 (Ancylis); Proc. En- 
tomol. Soc. Wash. 6: 221. Eight syn- 
types, USA, Rhode Island, Washington 
Co., Weekapaug, r.f. Lathyrus maritima, 
H. Dyar. One of the syntypes was 
placed in the type collection, presum- 
ably by J. Clarke; the remainder are in 
the main collection. 

marloffiana Busck, 1907 (Phalonia); J. N. 
Y. Entomol. Soc. 15: 26. Holotype °, 
USA, Pennsylvania, Allegheny Co., 
Oak Station, 7 June 1906, E Marloff. 

marmoreana Heinrich, 1923 (Epinotia); 
Bull. U.S. Natl. Mus. 123: 222. Holo- 
type d, USA, Utah, Tooele Co., Stock- 
ton, 16 July 1913, T. Spalding. 

marquesana Clarke, 1986 (Duessa); Smith- 
sonian Contrib. Zool. 416: 155. Holo- 
type 6, Marquesas Islands, Nuku Hiva, 
Tunoa Ridge, 2,900’, 23 January 1968, 
J. & T. Clarke. 

maurodicha Clarke, 1976 (Herpystis); In- 
sects of Micronesia 9: 74. Holotype 6, 
Micronesia, Kusaie, Mutunlik, 22 m, 21 
April 1953, J. Clarke. 

maximana Barnes & Busck, 1920 (Pero- 
nea), Contrib. Nat. Hist. Lepid. North 
Am. 4: 216. Holotype 6, Canada, Brit- 
ish Columbia, Victoria, A. Croker. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


mayelisana Blanchard, 1979 (Phaneta); J. 
Lepid. Soc. 33: 209. Holotype 6, USA, 
Texas, Cottle Co., Paducah, 17 April 
1968, A. & M. Blanchard. 

medioalbana Knudson, 1986 (Pammene); J. 
Lepid. Soc. 40: 325. Holotype ¢, USA, 
Texas, Gonzales Co., Palmetto State 
Park, 23 March 1985, E. Knudson. 

mediopartitum Heinrich, 1923 (Exartema); 
Proc. Entomol. Soc. Wash. 25: 113. Ho- 
lotype ¢, USA, Virginia, 1 June 1884. 

megalorhis Diakonoff, 1982 (Sychnochlae- 
na); Zool. Verhandel. (Leiden) 193: 
117. Holotype 2, Sri Lanka, NE Dis- 
trict, Kanda-ele Reservoir, 5.6 mi NW 
Nuwara Eliya, 6,200’, 10—21 February 
1970, D. Davis & B. Rowe. 

megasaccula Brown, 1991 (Bidorpitia); in 
Brown and Powell, Univ. Calif. Publ. 
Entomol. 111: 73. Holotype ¢, Guate- 
mala, Purulha, July, Schaus & Barnes. 

melanantha Diakonoff, 1968 (Peridaeda- 
la); Bull. U.S. Natl. Mus: 257: 77 sHe- 
lotype 2, Philippine Islands, Mindanao, 
Davao Prov., E slope Mt. McKinley, 
mossy stunted forest, at light, 19 Sep- 
tember 1946, CNHM Philippine Zool., 
Exped. 1946—47, H. Hoogstraal & D. 
Heyneman. 

melania Clarke, 1955 (Orthocomotis); 
Trans. Royal Entomol. Soc. London 
107: 153. Holotype 6, Jamaica, St. Ann 
Parish, 1,750’, E. Bell. 

melanomesum Heinrich, 1923 (Exartema); 
Proc. Entomol. Soc. Wash. 25: 119. Ho- 
lotype 6, USA, Maine, Piscataquis Co., 
Sebec Lake, “July 16—23.” 

melanoleuca Clarke, 1968 (Irazona); Proc. 
U.S. Natl. Mus. 125: 44. Holotype ¢d, 
Mexico, Puebla, rf. Pinus, January 
1960, Guevara. 

melasma Clarke, 1968 (Hysterosia); Proc. 
U.S. Natl. Mus. 125: 9. Holotype 4, 
Guatemala, Chejel, June, Schaus and 
Barnes. 

melia Clarke, 1976 (Adoxophyes); Insects 
of Micronesia 9: 133. Holotype 6, Mi- 
cronesia, Guam, Fadian, 19 August 
1936, ex-Colubrina, O. Swezey. 


VOLUME 102, NUMBER 4 


melidora Razowski 1984 (Ardeutica); Acta 
Zool. Cracov. 27: 217. Holotype 4, 
Cuba, Sierra del Cobra Oriente, Loma 
del Gato, 2,600’, 24-30 September 
1935, J. Acufia, S. Bruner & L. Scara- 
muzza. 

meligma Clarke, 1986 (Dichelopa); Smith- 
sonian Contrib. Zool. 416: 141. Holo- 
type 6, Marquesas Islands, Fatu Hiva, 
Tahuna, 2,000’, 22 March 1968, J. & T. 
Clarke. 

membrosa Heinrich, 1926 (Laspeyresia); 
Bull. U.S. Natl. Mus. 132: 49. Holotype 
2, USA, Texas, Bexar Co., San Anto- 
Mower. w_rosopis.s 290 June, 1917, A- 
Busck. 

mendaciana Blanchard & Knudson, 1983 
(Suleima); Proc. Entomol. Soc. Wash. 
85: 848. Holotype 6, USA, Texas, 
Brewster Co., Big Bend National Park, 
Dugout Wells, 28 September 1981, E. 
Knudson. 

mendora Clarke, 1968 (Cochylis); Proc. 
U.S. Natl. Mus. 125: 24. Holotype d, 
Chile, Santiago Province, Cajon de 
Maypo, Cordillera, El Canelo, 12-20 
January 1948, T. Ramirez. 

mengelana Fernald, 1894 (Sericoris); En- 
tomol. News 5: 131: Lectotype ¢, 
Greenland, McCormick Bay. Designat- 
ed by Miller (1970). 

meridionalis Yasuda & Kawabe, 1980 (Ar- 
chips); Tinea 11: 9. Holotype 6, Japan, 
Hatsuno, Is. Amami-oshima, 11—13 Au- 
gust 1977, A. Seino. 

meritana Heinrich, 1923 (Epinotia); Bull. 
U.S. Natl. Mus 123: 226. Holotype ¢, 
USA, Utah, Carbon Co., Hiawatha, rf. 
Pinus em: July 1921, H.<Peck. 

messalina Clarke, 1971 (Dichelopa),; 
Smithson. Contrib. Zool. 56: 102. Ho- 
lotype 2, Rapa Island, Maugaoa, 950° 
@92sm)e7 Nov 1963.5. &. 1. Clarke: 

metallica Busck, 1914 (Evetria); Proc. En- 
tomol. Soc. Wash. 16: 146. Holotype 6, 
USA, Montana, Missoula Co., Missou- 
la, r.f. pitch of yellow pine, 2 May 1913, 
J. Brunner. 

metariana Heinrich, 1923 (Eucosma); Bull. 


1047 


U.S. Natl. Mus. 123: 133. Holotype 6, 
USA, California, Siskiyou Co., Shasta 
Retreat, “June 16—23.” 

mexicana Busck, 1907 (Pharmacis); J. N. 
Y. Entomol. Soc. 15: 30. Holotype ¢, 
USA, New Mexico, Beulah, 8,000’, T. 
Cockerell, USNM type 10238. 

mezion Razowski 1984 (Ardeutica); Acta 
Zool. Cracoye, 27; 4218, Holotype 6: 
Cuba, Sierra del Cobre Oriente, Loma 
del Gato, 2,600’, 24-30 September 
1935, J. Acuna, S. Bruner & L. Scara- 
muzza. 

mesostigmatias Diakonoff, 1977 (Thauma- 
tographa),; Zool. Verhandel. (Leiden) 
158: 42. Holotype 6, Taiwan, Rantais- 
an, 15 May 1933;,S. Issiki: 

messalina Clarke, 1971 (Dichelopa); 
Smithsonian Contrib. Zool. 56:102. Ho- 
lotype 2, Rapa Island, Maugaoa, 950’ 
[292 m], 7 November 1963, J. & T. 
Clarke. 

microptera Clarke, 1953 (Endothenia),; J. 
Wash. Acad. Sci. 43: 230. Holotype 6, 
USA, Illinois, Putnam Co., 24 August 
1949, M. Glenn. 

mieae Kawabe, 1980 (Lobesia); Tinea 11: 
20. Holotype 6, Japan, Narahara, Hach- 
iooji-shi, Tokyo, 9 August 1978, A. Ka- 
wabe. 

migratana Heinrich, 1923 (Thiodia); Bull. 
U.S. Natl. Mus. 123: 53. Holotype 4, 
USA, California, Inyo Co., Olanche, 
“Apr 24-30.” 

mimica Clarke, 1976 (Herpystis); Insects of 
Micronesia 9: 78. Holotype °, Micro- 
nesia, Palau Islands, Koror Island, Ko- 
ror, 29 May 1957, C. Sabrosky. 

minor Brown 1990 (Macrochlidia); J. N. Y. 
Entomol. Soc. 98: 373. Holotype ¢, 
Venezuela, Aragua, Charoni Pass, 1,400 
m, 20 January 1965, S. & W. Duck- 
worth. 

minorata Heinrich, 1924 (Eucosma gigan- 
teana); J. Wash. Acad. Sci. 14: 388. Ho- 
lotype ¢, USA, Texas, Liberty Co., Lib- 
erty, em: 28 July 1922, FE Bottimer. 

minutana Kearfott, 1905 (Eucosma); Proc. 
U.S. Natl. Mus. 28: 356. Lectotype 6, 


1048 


USA, New Jersey, Essex Co., Montclair, 
July 1908, W. Kearfott. Designated by 
Blanchard (1979). 

mira Heinrich, 1929 (Anchylopera); Proc. 
U.S. Natl. Mus. 75: 17. Holotype 6, 
USA, Colorado, Jefferson Co., Chim- 
ney Gulch, Golden, April, Oslar. 

mira Razowski, 1989 (Saphenista); SHI- 
LAP Revista Lepidopterologia 17: 206. 
Holotype 2, Guatemala, Volcan Santa 
Maria, November, Schaus & Barnes. 

miramundi Razowski, 1988 (Clarkenia); 
Acta Zool. Cracov. 31: 407. Holotype 
?, El Salvador, Cerro Miramundo, 233 
January 1971, S. Steinhauser. 

mirosignata Heinrich, 1929 (Eucosma); 
Proc. U.S. Natl. Mus. 75: 11. Holotype 
3, USA, Arizona, Pima Co., Baboqui- 
vari Mountains, 15-30 October 1924, 
O. Poling. 

miscitata Heinrich, 1926 (Laspeyresia); 
Bull. U.S. Natl. Mus. 132: 64. Holotype 
3, USA, California, Shasta National 
Forest, rf. Pinus jeffreyi, May 1912, 
Hopk. 11414, J. Miller. 

misturana Heinrich, 1923 (Thiodia); Bull. 
U.S. Natl. Mus. 123: 54. Holotype 6, 
Canada, Saskatchewan, Oxbow, 9 June 
1907, E Knab. 

mobilensis Heinrich, 1923 (Eucosma); Bull. 
U.S. Natl. Mus. 123: 125. Holotype ¢, 
USA, Alabama, Baldwin Co., near 
Daphne, eastern shore of Mobile Bay, 
‘from root cuttings of Chrysoma (Sol- 
idago) pauciflosculosa ... during Sep- 
tember (1920),”’ em: 3 October 1920, T. 
van Aller. 

mochana Busck, 1913 (Olethreutes); Insect. 
Inscit. Menst. 1: 92. Holotype @, British 
Guiana, Mocha, “‘bred from Vochysea 
guianensis,’’ H. Moore. 

modestana Busck, 1907 (Hysterosia); J. N. 
Y. Entomol. Soc. 15: 32. Holotype <4, 
USA, Pennsylvania, Allegheny Co., 
Pittsburgh, 2 July 1905, H. Engel. 

moffatiana Fernald, 1905 (Proteoteras); 
Canad. Entomol. 37: 16. Lectotype <6, 
Canada, Ontario, London. Designated 
by Miller (1970). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


molesta Busck, 1916 (Laspeyresia); J. 
Agric. Res. U.S; Dept: Agric. 77°373: 
Holotype 6, USA, Virginia, Arlington 
Co., Arlington, rf. peach, em: 12 Oc- 
tober 1916. 

molybdaina Clarke, 1976 (Adoxophyes); In- 
sects of Micronesia 9: 138. Holotype 6, 
Micronesia, Ponape, Colonia, 7 January 
19535 J:<Glarke: 

molybdina Clarke, 1968 (Carolella); Proc. 
U.S. Natl. Mus. 125: 53. Holotype 2, 
Mexico, San Luis Potosi, Tamazun- 
chale, 6 January 1947, Chapman. 

monela Clarke, 1976 (Icelita); Insects of 
Micronesia 9: 39. Holotype 2, Micro- 
nesia, Kusaie, Mutunlik, 22 m, 29 Jan- 
uary 1953, J. Clarke. 

monetiferanum Riley, 1881 (Exartema); 
Trans. St. Louis Acad. Sci. 4: 317. Ho- 
lotype 6, USA, Alabama, Barbour Co., 
Eufala, G. Latimer. Riley described this 
species from a single specimen. 

monitorana Heinrich, 1920 (Eucosma); 
Proc. U.S. Natl. Mus. 57: 58. Holotype 
3, USA, Pennsylvania, Danville, rf. 
cones of Pinus, em: 4 May 1916, A. 
Champlain. 

monstrata Razowski, 1984 (Aphalonia); 
Ann. Zool. Warsz. 38: 276. Holotype 6, 
Peru, Divisoria, 5,200’, 20-23 June 
1982, C. Covell. 

montana Busck, 1914 (Evetria); Proc. En- 
tomol. Soc. Wash. 16: 147. Holotype 3, 
USA. Montana, Elliston, rf. Pinus con- 
torta, em: 10 December 1913, J. Brun- 
ner. 

montana Bartlett-Calvert, 1892 (Antithes- 
ia); Univ. Santiago Chile, Univ. Ann. 
84: 831. Neotype 6, Argentina, Neu- 
quén, Chapelco, Techos, 1,400 m, 24 
January 1984, M. & P. Gentili. Desig- 
nated by Brown (1998). 

monticolana Kawabe, 1964 (Clepsis); Tyo 
to Ga 15: 1. Holotype ¢, Japan, Toyama 
Pref., Honshu, Mt. Tateyama, 29 August 
1962, A. Kawabe. 

moriutit Kawabe, 1987 (Stenodes); Micro- 
lepid. Thailand 1: 72. Holotype 6, 
Thailand, Chiang Mai, Doi Pui, ca. 


VOLUME 102, NUMBER 4 


1,300 m, 26—27 October 1985, S. Mor- 
iuti, T. Saito & Y. Arita. 

mormonensis Heinrich, 1923 (Evetria); 
Bull. U.S. Natl. Mus. 123: 44. Holotype 
36, USA, Utah, Sal Lake Co., Salt Lake 
City, C. Ainslie. 

murtfeldtiana Riley, 1881 (Anchylopera); 
Trans. St. Louis Acad. Sci. 4: 323. 
“Type’’ ¢, USA, Missouri, rf. oak, em: 
19 May 1875, M. Murtfeldt. Riley de- 
scribed this species from three speci- 
mens, only one of which could be lo- 
cated by us. 

musetta Blanchard & Knudson, 1983 (Pha- 
neta); Proc. Entomol. Soc. Wash. 85: 
845. Holotype 6, USA, New Mexico, 
Socorro Co., Gran Quivara Natl. Mon- 
ument, 6,600’, 1-3 July 1964, D. Davis. 

mydros Obraztsov, 1966 (Idolatteria); Proc. 
U.S. Natl. Mus. 119: 6. Holotype ¢, Ec- 
uador, Environs de Loja, 1887. 

myopori Clarke, 1971 (Dichelopa); Smith- 
sonian Contrib. Zool. 56: 115. Holotype 
6, Rapa Island, Pariati Bay, rf. Myopo- 
rum rapensis, em: 30 Nov 1963, J. & T. 
Clarke. 

myoxa Razowski, 1984 (Polyortha); Acta 
Zool. Cracov. 27: 220. Holotype ¢, 
Brazil, Santa Catarina, Nova Teutonia, 
November 1961, E Plaumann. 


naevifera Razowski, 1984 (Polyortha); 
Acta Zool. Cracov. 27: 223. Holotype 
6, Venezuela, Aragua, Rancho Grande, 
10-21 February 1969, Duckworth & 
Dietz. 

nakajimai Kawabe, 1992 (Acleris); Tinea 
13: 172. Holotype 6, Taiwan, Hualien 
Hsien, Houhuanshan, Luoying Lodge, 
2,800 m, 31 December 1988, A. Ka- 
wabe. 

naoma Clarke, 1953 (Epiblema); J. Wash. 
Acad. Sci. 43: 226. Holotype ¢, USA, 
Illinois, Putnam Co., rf. Ratibida pin- 
nata, 25 June 1950, M. Glenn. 

nebulosana Packard, 1867 (Grapholitha); 
Proc. Boston Soc. Nat. Hist. 11: 61. 
Syntype 6, Canada, Labrador, Straw- 
berry Harbor, 26—30 July. Although the 


1049 


original description does not indicate 
how many specimens were examined 
nor identify a type, it states that the spe- 
cies “is not infrequent at Strawberry 
Harbor ...” 

negligens Kawabe, 1978 (Neoanathamna); 
Tinea 10: 185. Holotype 6, Japan, Shi- 
koku, Kagawa Pref., Mt. Zozu, 22 May 
1968, H. Toshima. 

negundana Dyar, 1902 (Cacoecia); in Cau- 
dell, Proc. Entomol. Soc. Wash. 5: 78. 
“Type” 6, USA, Colorado, El Paso 
Co., Pike’s Peak, Dyar & Caudell. Dyar 
briefly diagnosed this species in a report 
by Caudell (1902) without mentioning 
any specimens or designating a type. 

neoclyta Razowski, 1988 (Clarkeulia); Acta 
Zool. Cracov. 31: 405. Holotype 6, Co- 
lombia, Narino, Volcan Galeras, 2,900 
m, 13 January 1959, J. Clarke. 

neomexicana Dyar, 1903 (Evetria); Proc. 
Entomol. Soc. Wash. 5: 286. Holotype 
3, USA, New Mexico, San Miguel Co., 
Las Vegas. This species was described 
from the single male cited above. 

nepotinana Heinrich, 1923 (Thiodia); Bull. 
U.S. Natl. Mus. 123: 263. Holotype d, 
USA, Utah, Juab Co., Eureka, 30 May 
1911, T. Spalding. 

nephelodes Clarke, 1968 (Amallectis); Proc. 
U.S. Natl. Mus. 125: 27. Holotype ¢, 
Bolivia, Cochabamba, Incachaca, tropi- 
cal cloud area, 2,100 m, 27 August—5 
September 1956, L. Pena. 

nigra Miller, 1966 (Laspeyresia); J. Lepid. 
Socs ;20:0251. Holotype sd.Mexico, 
Tlaxcala, February 1964, “en semilla P. 
ayacahuite.”’ 

nigralbana Walsingham, 1879 (Paedisca); 
Ill. Lepid. Heter. Brit. Mus. 4: 41. Syn- 
type 6, USA, California, Mendocina 
Co., 10 June 1871. This species was de- 
scribed from two males and two fe- 
males; the other syntypes are presumed 
to be in the BMNH. 

nigricolor Yasuda & Kawabe, 1980 (Ka- 
wabeia); Tinea 11: 10. Holotype 6, Ja- 
pan, Nagano Pref. Azumi V., Shimaji- 


1050 


ma-dani, 780 m, 23 October 1976, N. 
Hirano. 

nigrilineana Kawabe, 1963 (Acleris); Tyo 
to Ga 14: 71. Holotype 6, Japan, Na- 
gano Pref., Honshu, Asama-sanso, 4 
May 1962, A. Kawabe. 

nigriplagana Franclemont, 1986 (Archips); 
Proc. Entomol. Soc. Wash. 88: 59. Ho- 
lotype ¢, USA, New York, Tompkins 
Co., McLean Bogs Reserve, 2 July 
1953, J. Franclemont. 

nigrivelata Walsingham, 1914 (Tortrix); 
Biol. Centr.-Am., Lepid., Heter. 4: 283. 
Holotype ¢, Panama, Canal Zone, Ta- 
bernilla, A. Busck. Walsingham de- 
scribed this species from a single male. 

nigromaculata Issiki, 1930 (Simaethis); 
Ann. Mag. Natl. Hist. 6: 423. Holotype 
?, Japan, Honsyu, Iwawakisan, 9 Au- 
gust 1918, S. Issiki. 

nigropunctata Kawabe, 1985 (Geogepa); 
Tinea 12: 7. Holotype d, Taiwan, Chiai 
Hsien, Fenchihu, 1,400 m, 8 August 
1971, Y. Shabata. 

ninana Dyar, 1903 (Carpocapsa); List Lep- 
id. North Am.: 471. Two syntypes (1 ¢, 
1 2), USA, Arizona, Fort Grant, 18 July 
1892. Authorship of this taxon (as Gra- 
pholitha ninana) usually is attributed to 
Riley; the name first appeared in Riley’s 
contribution to Smith’s (1891: 93) 
checklist of the Lepidoptera of North 
America without a description or refer- 
ence to specimens examined. Heinrich 
(1926: 60) correctly recognized that 
Dyar presented the first description, 
based on Riley’s specimens, in a foot- 
note in his checklist (Dyar 1903), and 
hence should be considered the author. 

nipponana Razowski, 1977 (Stenodes); Tyo 
to Ga 28: 35. Holotype 6, Japan, Si- 
koku, Asizuri-Saki, 10 May 1951, S. Is- 
siki. 

nipponica Kawabe, 1976 (Eucosma); Tinea 
11: 40. Holotype 6, Japan, Kanagawa 
Pref., Oyama, 13 May 1958, A. Kawa- 
be. 

nitida Clarke, 1955 (Orthocomotis); Trans. 
Royal Entomol. Soc. London 107: 143. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Holotype ¢d, Guatemala, Cayuga, “4,” 
Schaus & Barnes. 

nitida Horak, 1984 (Heterochorista); Syst. 
Entomol. 9: 429. Holotype 3, Papua 
New Guinea, Morobe District, Wau, 
1,200 m, mercury vapor light, 8-14 De- 
cember 1976, G. Hevel & R. Dietz. 

niveopunctata Dognin, 1905 (Polyortha); 
Ann. Soc. Entomol. Belgium 49: 85. 
Holotype 2, Ecuador, Environs de Loja, 
1893. 

niveosana Packard, 1867 (Sciaphila); Proc. 
Boston Soc. Nat. Hist. 11: 55. Holotype 
3d, Canada, Labrador, Hopedale, 3 Au- 
gust. The original description appears to 
have been based on a single male. 

nota Kawabe, 1978 (Cryptophlebia); Tinea 
10: 188. Holotype ¢, Japan, Honshu, 
Ishikawa Pref., Anamizu, Bira, 9 Au- 
gust 1961, A. Kawabe. 

notialis Miller, 1985 (Eucosma); Ann. En- 
tomol. Soc. Am. 78: 244. Holotype 6, 
USA, Texas, Kerr Co., Kerrville, 6 
June, Lacey. 

novimundi Heinrich, 1920 (Laspeyresia); 
Canad. Entomol. 52: 257. Holotype 3, 
USA, Wisconsin, Door Co., Sturgeon 
Bay, 14 July 1920, C. Fluke. 

nymphana Blanchard & Knudson, 1983 
(Gretchena); Proc. Entomol. Soc. Wash. 
85: 851. Holotype 6, USA, Texas, An- 
derson Co., Tennessee Colony, Gus En- 
geling Wildlife Management Area, 15 
April 1968, A. & M. E. Blanchard. 

nuntia Heinrich, 1929 (Eucosma); Proc. 
U.S. Natl. Mus. 75: 10. Holotype 3, 
USA, Utah, Juab Co., Callao, 17 April 
1922: 

nythobia Clarke, 1971 (Cryptophlebia); 
Smithsonian Contrib. Zool. 56: 133. 
Holotype 2, Rapa Island, Haurei, rf. 
Myoporum rapensis, 19 September 
1963, J. & T. Clarke. 


oachranta Diakonoff, 1974 (Polylopha); 
Zool. Verhandel. (Leiden) 131: 72. Ho- 
lotype ¢, Saipan Island, Marianas, 0.5 
mi E Tanapag, 20 April 1945, H. Dybas. 
obfuscana Riley, 1888 (Paedisca); Proc. 


VOLUME 102, NUMBER 4 


Entomol. Soc. Wash. 1: 33. ““Type” 3, 
USA, rf. Solidago, 2 June 1884. Riley 
did not indicate the number of speci- 
mens examined. 

obliqua Diakonoff, 1982 (Grapholita); 
Zool. Verhandel. (Leiden) 193: 20. Ho- 
lotype 6, Sri Lanka, Anuradhapura Dis- 
trict, Wildlife Sanctuary Bungalow, Hu- 
nuwilagama, Wilpattu, 200’, 10-19 
March 1970, Davis & Rowe. 

obnigrana Heinrich, 1923 (Proteoteras); 
Bull. U.S. Natl. Mus. 123: 169. Holo- 
type 6, USA, New Hampshire, Chesh- 
ire Co., Dublin, A. Busck. 

obnisa Heinrich, 1926 (Laspeyresia); Bull. 
U.S. Natl. Mus. 132: 47. Holotype °&, 
Canada, British Columbia, Fraser Mills, 
16 June 1922, E. Blackmore. 

occidentalis Heinrich, 1923 (Thiodia stria- 
tana); Bull. U.S. Natl. Mus. 123: 58. 
Holotype 6, USA, California, Siskiyou 
Co., Shasta Retreat, “‘July 16—23.” 

oceani Diakonoff, 1956 (Bactra); Zool. 
Verhandel. (Leiden) 29: 59. Holotype 
6, Fiji Islands, Nadi, at light, July, J. 
Tllinoworth. 

ochracea Clarke, 1955 (Orthocomotis); 
Trans. Royal Entomol. Soc. London 
107: 144. Holotype d6, Costa Rica, Car- 
tago Province, Juan Vinas, W. Schaus. 

ochraceana Fernald, 1901 (Epiblema); J.N. 
Y. Entomol. Soc. 9: 51. Holotype 4, 
USA, Florida, Palm Beach Co., Palm 
Beach. This species was described from 
a single male. 

ochroma Clarke, 1986 (Dichelopa); Smith- 
sonian Contrib. Zool. 416: 138. Holo- 
type ¢, Marquesas Islands, Nuku Hiva, 
Tapuaooa, 2,500’, 30 January 1968, J. 
& T. Clarke. 

ochrosaphes Clarke, 1955 (Orthocomotis); 
Trans. Royal Entomol. Soc. London 
107: 152. Holotype 6, Brazil, Santa Ca- 
tarina, 27 December 1935, EK Hoffman. 

ocliferia Heinrich, 1926 (Hemimene), Bull. 
U.S. Natl. Mus. 132: 21. Holotype 6, 
USA, Nevada, Washoe Co., Pyramid 
Lake. 

oenotherana 1881 


Riley, (Conchylis); 


1051 


Trans. St. Louis Acad. Sci. 4: 316. 
“Type”? 2, USA, Missouri, r.f. Oenoth- 
era, M. Murtfeld. Riley refers to four 
specimens in the original description; 
we have been able to locate only the 
specimen cited above. 

ogasawaraensis Kawabe & Ku, 1978 (Me- 
tendothenia); Tinea 10: 169. Holotype 
36, Japan, Ogasawara Island, Omura, 
Chichijima, 14 June 1972, Y. Watanabe. 

okubiensis Kawabe, 1987 (Eucosma); Tinea 
12: 141. Holotype ¢, Japan, Okinawa 
Island, Okubi, Kin, r.f. Bruguiera gym- 
norrhiza, 26 April 1983, S. Hori. 

olivaceana Fernald, 1882 (Eccopsis); Trans. 
Am. Entomol. Soc. 10: 71. Lectotype 
3, USA, Pennsylvania. Designated by 
Miller (1970). 

olivaceana Riley, 1881 (Grapholitha); 
Trans. St. Louis Acad. Sci. 4: 320. Ho- 
lotype ¢, USA, Illinois, J. Muhleman. 
Riley described this species from a sin- 
gle specimen. 

olivata Dognin, 1905 (Tortrix); Ann. Soc. 
Entomol. Belgium 49: 85. Holotype <6, 
Colombia, Popayan, 1895. 

onychistica Diakonoff, 1982 (Age); Zool. 
Verhandel. (Leiden) 193: 57. Holotype 
6, Sri Lanka, Mannar District, Mannar 
Island, 2 mi. NE Mannar, 15’, 22 March 
1970, D. Davis & B. Rowe. 

opistocapna Diakonoff, 1977 (Thaumato- 
grapha); Zool. Verhandel. (Leiden) 158: 
38. Holotype 6, New Guinea, Wareng, 
12 July 1936, S. Issiki. 

opposita Heinrich, 1931 (Epinotia); Proc. 
U.S. Natl. Mus. 79: 7. Holotype 4, 
Peru, Lima, D. Willie, ““No. 154.29.” 

optimana Dyar, 1893 (Eucosma),; Proc. En- 
tomol. Soc. Wash. 5: 180. Syntype ¢d, 
USA, Colorado, Garfield Co., Glen- 
wood Springs, ““Oct 1—7,’’ W. Barnes; 
syntype 2, USA, Colorado, Eagle Co., 
T. Cockerell. The two specimens listed 
comprise the type series. 

orcula Razowski, 1988 (Oregocerata); 
Acta Zool. Cracov. 31: 393. Holotype 
3, Bolivia, Cochabamba, Incachaca, 


1052 


tropical cloud area, 2,100 m, 27 Au- 
gust—5 September 1956, L. Pena. 

oregonensis Heinrich, 1923 (Thiodia); Bull. 
U.S. Natl. Mus. 123: 47. Holotype <4, 
USA, Oregon, Klamath Co., Crater 
Lake, “July 24-31.” 

oribasus Razowski, 1988 (Oryguncus); 
Acta Zool. Cracov. 31: 402. Holotype 
3, Peru, Cusco, Machu Picchu, 2,700 
m, 6 February 1959, J. Clarke. 

ornatula Heinrich, 1924 (Thiodia); J. Wash. 
Acad. Sci. 14: 385. Holotype 6, USA, 
Pennsylvania, Allegheny Co., Oak Sta- 
tion, 21 June 1909, FE Marloff. 

orphnoxantha Obraztsov, 1966 (Pseudo- 
meritastis); Proc. U.S. Natl. Mus. 118: 
227. Holotype 6, Costa Rica, Cartago 
Province, Tuis, 2,400’, W. Schaus. 

osmundana Fernald, 1879 (Penthina); Can- 
ad. Entomol. 11: 156. Lectotype °&, 
USA, Maine, Penobscot Co., Orono, 
July, r.f. O. regalis. Designated by Mill- 
er (1970). 

ostentatrix Heinrich, 1923 (Hystricophora); 
Bull. U.S. Natl. Mus. 123: 270. Holo- 
type 6, USA, California, Tulare Co., 
Mineral King, “‘Aug. 1-7.” Heinrich 
(1923) consistently misspelled HAystri- 
chophora in his revision. 

oxybela Razowski, 1988 (Ptynognathosia); 
Acta Zool. Cracov. 31: 394. Holotype 
3, Colombia, Cauca, Paramo de Parace, 
Lake San Rafael, 3,570 m, 29 January 
1959, J. Clarke. 

oxycoccana Packard, 1869 (Peronea); 
Guide to the Study of Insects: 334. 
“Type” 3d, USA, eastern states. This is 
the only Packard specimen of this spe- 
cies we could locate; it is clearly labeled 
as the type. Packard did not indicate 
how many specimens he examined. 

oxygona Diakonoff, 1968 (Eudemis); Bull. 
U.S. Natl. Mus. 257: 50. Holotype 2, 
Philippine Islands, Los Banfios, 2 July 
1932, A. Evangelista. 

oxyropa Razowski, 1990 (Ptoseulia); SHI- 
LAP Rev. Lepid. 18: 212. Holotype <4, 
Costa Rica, Cartago Province, Turrialba, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


22-28 February 1965, S. & W. Duck- 
worth. 

ozonia Razowski, 1990 (Ptoseulia); SHI- 
LAP Rev. Lepid. 18: 213. Holotype ?, 
Bolivia, Cochabamba, Incachaca, tropi- 
cal cloud area, 2,100 m, 27 August—5 
September 1956, L. Pefia. 


packardiana Fernald, 1886 (Tortrix); Bull. 
U.S. Dept. Agric. Entomol. 12: 20. Two 
syntypes (2 66), USA, Maine, Peaks 
Island, Casco Bay; and USA, New 
Hampshire; both specimens are labeled 
““cotype.’’ Freeman (1958) correctly in- 
dicated that the “‘type”’ is in the USNM, 
listing Peaks Island as the type locality. 
This could be interpreted as his desig- 
nation of the Maine specimen as the lec- 
toype. 

pallens Kawabe, 1980 (Neoanathamna); 
Tinea 11: 22. Holotype 6, Japan, Hi- 
totsu-uchi, Shionoe, Kagawa, 2 June 
1973, H. Tashima. 

pallidibasalis Heinrich, 1920 (Laspeyre- 
sia); Proc. U.S. Natl. Mus. 57: 60. Ho- 
lotype 6, USA, Oregon, Kaolin Beds, 
rf. cones of Abies concolor, 7 Septem- 
ber 1915, Sergent & Patterson. 

pallifasciata Powell, 1978 (Rhyacionia); in 
Powell and Miller, U.S. Dept. Agric., 
Agric. Handb. 514: 28. Holotype 6, 
USA, Arizona, Coconino Co., Fort Val- 
ley, 2.5 mi NW Flagstaff, 10 July 1981, 
R. Hodges. 

palmetum Heinrich, 1928 (Laspeyresia); 
Proc. Entomol. Soc. Wash. 30: 6. Ho- 
lotype 36, USA, Florida, near Royal 
Palms State Park, “‘II.4,’’ E Jones. 

palpana Walsingham, 1879 (Paedisca); Il. 
Lepid. Heter. Brit. Mus. 4: 54. Two syn- 
types (2 66), USA, California, Siski- 
you Co., Mount Shasta, August 1871. 
This species was described from five 
males and one female. All but the two 
syntypes listed above are presumed to 
be in the BMNH. 

paracinderella Powell, 1964 (Acleris); 
Univ. Calif. Publ. Entomol. 32: 96. Ho- 


VOLUME 102, NUMBER 4 


lotype 6, USA, California, Nevada Co., 
r.f., Prunus dumosa, C. Riley. 

paracornuta Brown, 1991 (Cuproxena); in 
Brown and Powell, Univ. Calif. Publ. 
Entomol. 111: 55. Holotype 6, Vene- 
zuela, Aragua, Rancho Grande, 1,100 
m, 16—23 October 1966, S. & W. Duck- 
worth. 

paradisiae Heinrich, 1923 (Hystricophora); 
Bull. U.S. Natl. Mus. 123: 255. Holo- 
type 6, USA, Washington, Pierce Co., 
Mount Rainier, Paradise Valley, “‘July 
24-30.” Heinrich (1923) consistently 
misspelled Hystrichophora in his revi- 
sion. 

paragnoma Clarke, 1986 (Dichelopa); 
Smithsonian Contrib. Zool. 416: 139. 
Holotype °, Marquesas Islands, Hiva 
Oa, Puamau, 8 February 1968, J. & T. 
Clarke. 

parallelana Walsingham, 1879 (Cochylis); 
Ill. Lepid. Heter. Brit. Mus. 4: 28. Syn- 
type 2, USA, California, Lake Co., 18 
June 1871. This species was described 
from two males and two females; the 
other syntypes are presumably in the 
BMNH. 

parana Busck, 1911 (Tortrix); Proc. U.S. 
Natl. Mus. 40: 228. Holotype 2, Brazil, 
Parana, Castro, “6234 Wlsm. 1908.” 

paraplesiana Blanchard, 1979 (Sonia); J. 
Lepid. Soc. 33: 184. Holotype ¢, USA, 
Texas, Harris Co., Houston, 5 June 
1968, A. & M. Blanchard. 

parmata Razowski, 1984 (Ardeutica); Acta 
Zool. Cracov. 27: 216. Holotype ¢, 
Costa Rica, Ojo de Agua, Rt. 2, km 75, 
20 July 1967, O. Flint. 

parva Brown, 1984 (Corticivora); Proc. 
Entomol. Soc. Wash. 86: 279. Holotype 
6, USA, Arkansas, Washington Co., 
Devil’s Den State Park, 24 June 1966, 
R. Hodges. 

parvana Kawabe, 1980 (Phalonidia); Tinea 
11: 29. Holotype 6, Japan, Yamanashi 
Pref., Kiyosato, 22 July 1972, A. Ka- 
wabe. 

passerana Walsingham, 1879 (Paedisca); 
Ill. Lepid. Heter. Brit. Mus. 4: 49. Syn- 


1053 


type 2, USA, California, Mendocino 
Co., 9-10 June 1871. According to the 
original description, this species was de- 
scribed from four males. Because the 
USNM specimen is almost certainly 
part of the type series, Walsingham ap- 
parently erred in his determination of 
the sexes. There is a congeneric male 
specimen in the type collection labeled 
as ‘““Paedisca plumigerana Wlsm. Cal. 
Type’’—apparently a Walsingham man- 
uscript name. 

pastigiata Heinrich, 1929 (Thiodia); Proc. 
U:S® Natl) Mus#i75: . 32 Holotyperd: 
USA, California, Tulare Co., Monachee 
Meadows, 8,000’, “July 8-14.” 

pediasios Miller, 1985 (Eucosma); Ann. 
Entomol. Soc. Am. 78: 241. Holotype 
3, USA, Texas, Terrel Co., Sanderson, 
25 April 1981, E. Knudson. 

penai Brown, 2000 (Lobogenesis); Proc. 
Entomol. Soc. Wash. 102: 27. Holotype 
36, Bolivia, Incachaca, Cochabamba, 
2,100 m, tropical cloud forest area, 27 
August—5 September 1956, L. Pefia. 

penai Clarke, 1968 (Amallectis); Proc. U.S. 
Natl. Mus. 125: 28. Holotype °, Boliv- 
ia, Cochabamba, Incachaca, tropical 
cloud area, 2,100 m, 27 August—5 Sep- 
tember 1956, L. Pena. 

penai Razowski, 1988 (nape); Acta Zool. 
Cracov. 31: 395. Holotype 6, Bolivia, 
Cochabamba, Incachaca, tropical cloud 
area, 2,100 m, 27 August-5 September 
1956, L. Pena. 

pennsylvaniana Kearfott, 1907 (Proteopte- 
ryx); Trans. Am. Entomol. Soc. 33: 48. 
‘Type’? 6, USA, Pennsylvania, Beaver 
Co., New Brighton, 22 March 1903, H. 
Merrick [genitalia slide only; we have 
been unable to find the pinned speci- 
men]. The original description of penn- 
sylvaniana consists of the final para- 
graph of the description of Proteopertyx 
albicapitana: “‘It is not impossible that 
fuller knowledge of the life histories of 
the western and eastern forms may 
prove them to be separate and distinct, 
in which case I would propose the name 


1054 


pennsylvaniana for the latter.’ Klots 
(1942) makes no mention of this spe- 
cies; i.e., presumably no Kearfott spec- 
imens are in the AMNH. 

pergandeana Fernald, 1905 (Eucosma); 
Canad. Entomol. 37: 399. Lectotype 6, 
USA, Virginia, 4 June 1882. Designated 
by Miller (1970). 

periculosana Heinrich, 1923 (Epiblema); 
Bull. U.S. Natl. Mus. 123: 268. Holo- 
type 6, Canada, British Columbia, 
Mount McLean, 5,500’, August, A. 
Hanham. 

peroneana Barnes & Busck, 1920 (Tortrix); 
Contrib. Nat. Hist. Lepid. North Am. 4: 
214. Holotype 6, USA, Arizona, Co- 
chise Co., Palmerlee [Huachuca Moun- 
tians]. 

perplexana Fernald, 1901 (Epiblema); J. N. 
Y. Entomol. Soc. 9: 51. Holotype ¢, 
USA, Florida, Palm Beach Co., Palm 
Beach, H. Dyar. 

perpropinqua Heinrich, 1929 (Eucosma); 
Proc. U.S. Natl. Mus. 75: 8. Holotype 
3d, USA, Arizona, Pima Co., Indian Oa- 
sis, Sells Post Office, 15-30 April 1923, 
O. Poling. 

persolita Heinrich, 1929 (Eucosma); Proc. 
U. S. Natl. Mus. 75: 7. Holotype ¢, 
USA, Texas, Cameron Co., San Benito, 
*“March 16—23.”’ 

perspicua Diakonoff, 1968 (Sycacantha in- 
odes); Bull. U.S. Natl. Mus. 257: 61. 
Holotype 6, Philippine Islands, Luzon, 
Mt. Makiling, Baker. 

perspicuana Barnes & Busck, 1920 (Hys- 
terosia); Contrib. Nat. Hist. Lepid. 
North Am. 4: 218. Holotype 2, USA, 
Arizona, Cochise Co., Paradise, August. 

peruvianus Razowski, 1988 (Telurips); 
Acta Zool. Cracov. 31: 391. Holotype 
3, Peru, Cusco, Machu Picchu, 2,700 
m, 6 February 1959, J. Clarke. 

phaeostropha Clarke, 1976 (Duessa); In- 
sects of Micronesia 9: 30. Holotype &, 
Micronesia, Rota, Rota, 22 m, at light, 
20 July 1946, Townes 805. 

phalaenopa Diakonoff, 1968 (Argyropto- 
cha); Bull. U.S. Natl. Mus. 257: 70. Ho- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


lotype ¢, Philippine Islands, Luzon, Mt. 
Makiling, Baker. 

pharetrata Razowski, 1987 (Terinebrica); 
Tinea 12(suppl.): 136. Holotype 3, Ar- 
gentina, Tucuman, Ciudad Universitar- 
ia, 800 m, 17 February 1959, J. Clarke. 

philerocherda Diakonoff, 1964 (Bactra); 
Zool. Verhandel. (Leiden) 70: 31. Ho- 
lotype ¢, British West Indies, Domini- 
ca, Antrim, 1,000’, 13 March 1956, J. 
Clarke. 

physoptila Diakonoff, 1968 (Peridaedala); 
Bull. U.S. Natl. Mus. 257: 75. Holotype 
6, Philippine Islands, Davao Province, 
Mindanao, Mt. Apo, 7,000’, at light, 
November 1946, CNHM Philippine 
Zool. Exped., Alcasid. 

piceae Issiki, 1961 (Panoplia); in Issiki & 
Mutuura, Shinyogu Kagai Shogarui: 36. 
Holotype ¢, Japan, Honsyu, Sinano, 
Omekura, 14 July 1957, rf. pupa on 
German Picea, T. Yasuda. 

piceaea Busck, 1916 (Olethreutes); Proc. 
Entomol. Soc. Wash. 18: 151. Holotype 
3d, USA, Colorado, El Paso Co., Colo- 
rado Springs, “bred during May and 
July in the Forest Insect Division from 
Picea parryana and Picea engelman- 
ni,’ J. Polloch & B. Harvey. 

picicola Dyar, 1906 (Eucosma); J. N. Y. En- 
tomol. Soc. 14: 108. Holotype 2, USA, 
Washington, Pierce Co., Mt. Rainier, 
Paradise Valley, rf. Abies lasiocarpa, 
H. Burke. According to the original de- 
scription, this species was described 
from a single female. 

pimana Busck, 1907 (Phalonia); J. N. Y. 
Entomol. Soc. 15: 24. Holotype 9°, 
USA, Arizona, Pima Co., Baboquivera 
Mtns., August. 

piperana Busck, 1906 (Hemimene); Proc. 
Biol. Soc. Wash. 19: 177. Holotype 6, 
USA, Washington, Whitman Co., Pull- 
man, 19 July 1898. 

pithecolobiae Busck, 1934 (Gymnandroso- 
ma); Entomol. Am. 13: 156. Holotype 
3d, Cuba, Havana Province, Santiago, 
Vegas, 15 July 1931, A. Otero. 

placidus Yasuda & Kawabe, 1980 (Acleris); 


VOLUME 102, NUMBER 4 


Tinea 11: 12. Holotype 6, Japan, Nip- 
para, Tokyo, 11 April 1964, A. Kawabe. 

platina Clarke, 1968 (Irazona); Proc. U.S. 
Natl. Mus. 125: 45. Holotype 6, Costa 
Rica, Alajuela Province, Mount Poas, 
May. 

platyxantha Clarke, 1986 (Dichelopa); 
Smithsonian Contrib. Zool. 416: 131. 
Holotype 2, Marquesas Islands, Fatu 
Hiva, Mt. Teoiaua, 2,000’, 22 March 
1968, J. & T. Clarke. 

pleodontia Razowski, 1987 (Monortha), 
Bull. Polish Acad. Sci. 35: 65. Holotype 
36, Panama, Rio Trinidad, March 1912, 
A. Busck. 

pleurogramma Clarke, 1976 (Duessa); In- 
sects of Micronesia 9: 28. Holotype 6, 
Micronesia, Kusaie, Mutunlik, 22 m, 30 
March 1953, J. Clarke. 

plicata Brown, 1992 (Catastega); J. N. Y. 
Entomol. Soc. 100: 224. Holotype 2°, 
USA, Arizona, Santa Cruz Co., Santa 
Rita Mountains, Madera Canyon, Bog 
Spring Camp Ground, 5,100’, 10-26 
July 1964, D. Davis. 

plumbata Clarke, 1951 (Anchylopera); J. 
Wash. Acad. Sci. 41: 296. Holotype ¢, 
Argentina, Tigre, May 1944, E Bour- 
quin. 

plumbosana Kawabe, 1972 (Olethreutes); 
Tinea 9: 244. Holotype ¢, Toyama 
Pref., Ohmaki, 3 June 1951, M. Hoshi- 
no. 

plummeriana Busck, 1906 (Hemimene); 
Proc. Biol. Soc. Wash. 19: 181. Holo- 
type 6, USA, Maryland, Montgomery 
Co., Plummers Island (in Potomac Riv- 
er above Washington, D.C.), May 1906. 

plummeriana Busck, 1907 (Phalonia); J. N. 
Y. Entomol. Soc. 15: 24. Holotype 6, 
USA, Maryland, Montgomery Co., 
Plummers Island, 1 May 1903, A. 
Busck. 

poecilogramma Clarke, 1976 (Adoxo- 
phyes); Insects of Micronesia 9: 125. 
Holotype 6, Micronesia, Kusaie, Hill 
1010, 300 m, light trap, 13 April 1953, 
J. Clarke. 

pollexifera Razowski, 1989 (Tamboma- 


1055 


chaya); SHILAP Revta. Lepid. 17: 205. 
Holotype 6, Peru, Cusco, Tamboma- 
chay 3,690 m, 3 February 1959, J. 
Clarke. 

polysticta Clarke, 1976 (Cryptaspasma); In- 
sects of Micronesia 9: 63. Holotype &, 
Micronesia, Kusaie, Mt. Matante, 300 
m, 1 February 1953, J. Clarke. 

ponderosa Powell, 1968 (Eucosma); Hil- 
gardia,, 39:13.) Holotype, (GasUSA. 
Oregon, Lake Co., Embody’s Mill, nr. 
Silver Lake, r.f. Pinus ponderosa, 5 Au- 
gust 1915,,P Sergent. 

ponderosana Obraztsov, 1962 (Choristo- 
neura lambertiana); Am. Mus. Novit. 
2101: 14. Holotype ¢, USA, Colorado, 
Sugarloaf, rf. Pinus ponderosa, 7 Au- 
gust 1937, J. Beal: 

ponera Walsingham, 1914 (Tortrix); Biol. 
Centr.-Am., Lepid., Heter. 4: 279. Ho- 
lotype 3d, Mexico, Puebla, Popocateptl 
Park, 13,000’, W. Schaus. Walsingham 
clearly identifies the USNM male as 
‘““Type”’ in the original description. 

poolei Brown, 1991 (Bidorpitia); in Brown 
and Powell, Univ. Calif. Publ. Entomol. 
111: 73. Holotype 6, Venezuela, Ara- 
gua, Rancho Grande, 1,100 m, 8—14 
August 1967, R. Poole. 

populana Busck, 1916 (Laspeyresia); Proc. 
Entomol. Soc. Wash. 18: 151. Holotype 
2, USA, Montana, Missoula Co., Mis- 
soula, 15 June 1914, J. Brunner. 

postica Bae, 1993 (Lobesia); Japan. J. En- 
tomol. 61: 522. Holotype 2, Taiwan, 
Pintung Hsien, Kenting Park, 100 m, 
26-27 March 1984, A. Kawabe. 

pottsi Clarke, 1976 (Dudua); Insects of Mi- 
cronesia 9: 94. Holotype 6, Micronesia, 
Truk, Wena (Moen), Civil Administra- 
tion Area, 6 March 1949, R. Potts. 

praesumptiosa Heinrich, 1923 (Epiblema); 
Bull. U.S. Natl. Mus. 123: 143. Holo- 
type 6, USA, Texas, Cameron Co., 
Brownsville, 27 May 1917, A. Busck. 

prochaldera Clarke, 1955 (Orthocomotis); 
Trans. Royal Entomol. Soc. London 
107: 145. Holotype ¢, Equador, Envi- 
rons de Loja, 1890. 


1056 


proxima Clarke, 1976 (Dudua); Insects of 
Micronesia 9: 91. Holotype ¢, Micro- 
nesia, Ponape, Colonia, 9 January 1953, 
J. Clarke. 

pruni Heinrich, 1923 (Anchylopera burges- 
siana); Bull. U.S. Natl. Mus. 123: 239. 
Holotype 6, USA, New Jersey, Cald- 
well, 17 May 1903, W. Kearfott. 

psathyra Diakonoff, 1989 (Lopharcha); 
Tinea 12: 202. Holotype 6, Japan, Kyu- 
syu, Hikosan, 15 July 1937, S. Issiki. 

pseudogamma Brown, 1990 (Paraptila); J. 
Lepid. Soc. 44: 265. Holotype 6, El 
Salvador, Santa Tecla, 28-29 October 
1967, E. Todd. 

pseudolivata Clarke, 1955 (Orthocomotis); 
Trans. Entomol. Soc. London 107: 147. 
Holotype 6, Brazil, New Bremen, 7 
October 1936, E Hoffman. 

pseudomalesana Clarke, 1986 (Cydia); 
Smithsonian Contrib. Zool. 416: 164. 
Holotype ¢, Marquesas Islands, Fatu 
Hiva, Omoa, 16 March 1968, J. & T. 
Clarke. 

pseudoplesia Brown, 1991 (Cuproxena); in 
Brown and Powell, Univ. Calif. Publ. 
Entomol. 111: 60. Holotype 3, Vene- 
zuela, Aragua, Rancho Grande, 1,100 
m, 22-31 July 1967, R. Poole. 

pseudotsugana Kearfott, 1904 (Cydia); 
Canad. Entomol. 36: 110. Holotype 6, 
Canada, British Columbia, Kaslo, rf. 
Pseudotsuga, larva collected 24 June. 
Although the specimen bearing the 
Kearfott type label (Type No. 7788) has 
no locality data, the pupal shell and as- 
sociated plant material indicate that it is 
the specimen referred to as ‘“‘also bred 
by him [Dyar] from larvae taken June 
24 on Pseudotsuga sp.” 

ptycta Razowski, 1987 (Popayanita); Tinea 
12(suppl.): 126. Holotype 6, Colombia, 
Cauca, 17 km SE Popayan, 2,000 m, 10 
January 1959, J. Clarke. 

ptygma Razowski, 1993 (Apotoforma); 
Acta Zool. Cracov. 36: 186. Holotype 
2, Mexico, San Luis Potosi, 8 August 
1966, O. Flint. Although the specimen 
is not labeled as the holotype, Razows- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ki’s description was clearly based on 
this single female specimen. 

pulchella Kawabe, 1963 (Acleris); Tyo to 
Ga 14: 71. Holotype 6, Japan, Honshu, 
Tokyo, Tama Hills, 27 June 1958, M. 
Suzuki. 

pulcheria Clarke, 1971 (Dichelopa); Smith- 
sonian Contrib. Zool. 56: 120. Holotype 
6, Rapa Island, Perau, 1,900’, 15 Oc- 
tober 1963, J. & T. Clarke. 

pulcherrima Razowski, 1971 (Acleris); 
Acta Zool. Cracov. 16: 548. Holotype 
?, Taiwan, Hassenzan, 5 June 1942, S. 
Issiki. 

pulsatillana Dyar, 1903 (Eucosma); Proc. 
Entomol. Soc. Wash. 5: 297. Nineteen 
syntypes, USA, Colorado, Jefferson 
Co., “‘foothills at Boulder and Golden,”’ 
rf. Pulsatilla hirsutissima. According to 
the original description, this species was 
described from 20 males and eight fe- 
males. 

pulveratana Walsingham, 1879 (Paedisca); 
Ill. Lepid. Heter. Brit. Mus. 4: 45. Syn- 
type 6, USA, California, San Francisco, 
16 May 1871. This species was de- 
scribed from four males, three of which 
are presumed to be in the BMNH. 

punctidiscanum Dyar, 1904 (Gymnandro- 
soma); Proc. Entomol. Soc. Wash. 6: 
60. Two syntypes (2 ¢¢), USA, Wash- 
ington, D.C., May, L. Howard; Penn- 
sylvania, New Brighton. The two spec- 
imens comprise the original type series. 

purpuratus Kawabe, 1965 (Archips); Tyo to 
Ga 16: 16. Holotype 3, Japan, Gunma 
Pref., Mt. Akagi, 19-21 July 1964, A. 
Kawabe. 

purpurea Dognin, 1904 (Atteria); Ann. Soc. 
Entomol. Belgium 48: 132. Holotype @, 
Ecuador, Environs de Loja, 1893. 


quadrifasciana Fernald, 1882 (Tortrix); 
Trans. Am. Entomol. Soc. 10: 67. Two 
syntypes (2 66), USA, Maine. Accord- 
ing to the original description, Fernald 
had specimens from Maine, New 
Hampshire, Massachusetts, New York, 
and Illinois. Although the number of 


VOLUME 102, NUMBER 4 


specimens examined was not given in 
the description, Fernald’s unpublished 
list of types donated to the USNM in- 
dicates three “‘cotypes.”’ 

quercana Fernald, 1882 (Cenopis); Trans. 
Am. Entomol. Soc. 10: 69. Lectotype ? 
(designated here), USA, Texas, Dallas 
Co., Dallas, 29 May. Selected by Powell 
in 1965. Fernald described this species 
from four males and four females from 
New York, Missouri, and Texas. 

quercifoliana Fitch, 1858 (Tortrix); New 
Work Agric: (Rept;,.826 “Type” «<, 
USA, New York [no locality data on 
specimen label]. Fitch did not indicate 
the number of specimens he examined; 
this is the only Fitch specimen of this 
species we could find. 


radicana Heinrich, 1923 (Griselda); Bull. 
U.S. Natl. Mus. 123: 186. Lectotype ¢, 
Canada, British Columbia, on spruce, 
G. Hewitt. Designated by Obraztsov 
(1965). 

rafaeliana Razowski, 1989 (Saphenista); 
SHILAP Revta. Lepid. 17: 206. Holo- 
type 6, Colombia, Cauca, Paramo de 
Parace, Lake San Rafael, 3,570 m, 27 
January 1959, J. Clarke. 

rana Busck, 1907 (Phalonia); J. N. Y. En- 
tomol. Soc. 15: 22. Holotype 6, USA, 
Pennsylvania, Allegheny Co., Pitts- 
burgh, 30 July 1905, H. Engel. 

razowskii Kawabe, 1963 (Tortricodes) Tin- 
ea 6: 7. Holotype 6, Japan, Chiba Pref., 
Narashino, 4 March 1958, A. Kawabe. 

razowskit Kawabe, 1992 (Clepsis); Tinea 
13: 178. Holotype 3, Taiwan, Hualien 
Hsien, Kuanyuan, 2,400 m, 7-8 August 
1987, A. Kawabe. 

reprobata Clarke, 1976 (Lobesia); Insects 
of Micronesia 9: 104. Holotype °, Mi- 
cronesia, Guam, Pt. Oca, 4 June 1945, 
Bohart and Gressitt. 

rescissoriana Heinrich, 1920 (Eucosma); 
Proc. U.S. Natl. Mus. 57: 58. Holotype 
3, USA, Oregon, Sprague River, rf. 
cones of Pinus murrayana, 30 May 
1914, P. Sergent. 


1057 


retrusa Razowski, 1993 (Acleris); Acta 
Zool. Cracov. 36: 191. Holotype °&, 
Mexico, Jalapa, November 1963, rf. 
Rubus, N. Krauss. 

rhodographa Clarke, 1971 (Dichelopa); 
Smithsonian Contrib. Zool. 56: 103. 
Holotype ¢, Rapa Island, Perau, 1000’ 
(307 m), rf. Veronica rapensis, em: 18 
November 1963, J. Clarke. 

rhoifructana Kearfott, 1904 (Polychrosis); 
Trans. Am. Entomol. Soc. 30: 296. Ho- 
lotype ¢, USA, Ohio, Wayne Co., Woo- 
ster, Insectory, 29 January 1884, Murt- 
feldt. 

rigidana Fernald, 1880 (Retinia); in Com- 
stock, Annu. Rep. Dept. Agric. 1880: 
237. Lectotype 2, USA, New York, 
Tompkins Co., Ithaca. Designated by 
Miller (1970). 

rosaocellana Knudson, 1986 (Eucosma); J. 
Lepid. Soc. 40: 322. Holotype 36, USA, 
Texas, Hemphill Co., Gene Howe 
WMA, 18 May 1985, E. Knudson. 

rubra Kawabe, 1992 (Neopotamia); Japan. 
Hetero. J. 170: 358. Holotype 3, Tai- 
wan, Huslien Hsien, Hohuanshan, 3,100 
m, 30 July—1 August 1983, A. Kawabe. 

ruidosana Heinrich, 1923 (Epinotia); Bull. 
U.S. Natl. Mus. 123: 216. Holotype 6, 
USA, New Mexico, Lincoln National 
Forest, Ruidosa Canyon, larvae mining 
leaves of “‘Alum root”? [Heuchera woo- 
toni], 8 May 1917, C. Heinrich. 

ruptimacula Dognin, 1904 (Tortrix); Ann. 
Soc. Entomol. Belgium 48: 132. Holo- 
type °, Ecuador, Cayanuma, Loja. 

russata Heinrich, 1924 (Epinotia cruciana); 
J. Wash. Acad. Sci. 14:391. Holotype 
3, Canada, British Columbia, Victoria, 
24 June 1923. 

russeola Heinrich, 1929 (Eucosma); Proc. 
U.S. Natl. Mus. 75: 6. Holotype 6, 
USA, California, Los Angeles Co., 11 
June 1921, K. Coolidge. 


sabiniana Kearfott, 1907 (Evetria); Trans. 
Am “Entomols'Soech 33:2. siypey ci, 
USA, California, Sacramento Co., rf. 
Pinus sabiniana, 9 August 1885. Kear- 


1058 


fott described this species from two 
specimens, one in the USNM collection 
and one in his personal collection. Al- 
though the Kearfott collection is depos- 
ited at the AMNH, Klots (1942) makes 
no mention of a specimen of this spe- 
cies there. 

saetigera Razowski, 1987 (Terinebrica); 
Tinea 12(suppl.): 137. Holotype 6, Bo- 
livia, Cochabamba, Incachaca, tropical 
cloud area, 2,100 m, 27 August—5 Sep- 
tember 1956, L. Pena. 

sagax Razowski, 1984 (Polyortha); Acta 
Zool. Cracov. 27: 221. Holotype ¢, Bo- 
livia, Cochabamba, Incachaca, J. Stein- 
bach. 

saileri Clarke, 1987 (Cryptophlebia); Acta 
Entomol. Chilean 14: 7. Holotype 6, 
Chile, Iquique Province, Region Tara- 
paca, Pampa Tamarugal, Fundo Refres- 
co, rf. Prosopis tamarugo, 29 June 
1985, D. Bobadilla. 

salaciana Blanchard & Knudson, 1981 (Eu- 
cosma); J. Lepid. Soc. 35: 176. Holo- 
type 6, USA, Texas, Nueces Co., North 
Padre Island, 13 October 1979, A. & M. 
Blanchard. 

saliciana Clemens, 1864 (Hedya); Proc. 
Entomol. Soc. Philadelphia 3: 515. Syn- 
type 36, USA, [probably eastern U.S.; 
no data on specimen label or in original 
description], rf. Salix longifolia, Walsh. 
Although Clemens indicated the num- 
ber of specimens examined and the de- 
pository (e.g., “Coll. Ent. Soc. Phila- 
delphia’’) for most species described in 
this work, he did not do so for this or 
the following species, H. salicicolana. 

salicicolana Clemens, 1864 (Hedya); Proc. 
Entomol. Soc. Philadelphia 3: 515. Two 
syntypes (1 6, 1 2), USA, [probably 
eastern U.S.; no data on specimen labels 
or in original description], r.f. Salix hu- 
milis, Walsh. 

salmicolorana Heinrich, 1923 (Thiodia); 
Bull. U.S. Natl. Mus. 123: 62. Holotype 
3, USA, Utah, Tooele Co., Stockton, 30 
July 1913, T. Spalding. 

salmonicolor Powell, 1978 (Rhyacionia); in 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Powell and Miller, U.S. Dept. Agric., 
Agric. Handb. 514: 15. Holotype 6, 
USA, Texas, Jeff Davis Co., Davis 
Mtns., Mt. Locke, 19 May 1971, A. & 
M. Blanchard. 

salvadorana Brown & Powell, 2000 (Ano- 
pina); Univ. Calif. Publ. Entomol. 120: 
57. Holotype 6, El Salvador, Cerro Mir- 
amundo, 2,300 m, 11 January 1969, S. 
Steinhauser. 

salweenensis Miller, 1977 (Petrova); J. 
Lepid. Soc. 31: 136. Holotype 6, Thai- 
land, Baw Luang, 20 March 1972, D. 
Chaiglom. 

sambuci Clarke, 1952 (Polychrosis); J. 
Wash. Acad. Sci. 43: 228. Holotype 36, 
USA, Illinois, Putnam Co., rf. Sambu- 
cus canadensis, 5 July, M. Glenn. 

santacrucis Obraztsov, 1963 (Acleris); 
Proc. U.S. Natl. Mus. 114: 225. Holo- 
type 6, USA, California, Santa Cruz 
Co., Santa Cruz, 29 October 1932, W. 
Tilden. 

santamariana Razowski, 1988 (Ortogna- 
thosia); Acta Zool. Cracov. 31: 392. 
Holotype ¢, Guatemala, Volcan Santa 
Maria, Barnes & Schaus. 

sapodilla Heppner, 1981 (Dichrorampha); 
Florida Entomol. 64: 271. Holotype 3d, 
USA, Florida, Dade Co., Homestead, 
rf. Achras zapota, 14 April 1974, R. 
Baronowski. 

saracana Kearfott, 1907 (Cenopis); Trans. 
Am. Entomol. Soc. 33: 68. “Type” 6, 
USA, New Jersey, Essex Co., Essex 
County Park, hills above Montclair, July 
1907, W. Kearfott. Kearfott described 
this species from one male and two fe- 
males, two field-collected and one 
reared from sassafras. Klots (1942) does 
not mention this species in his treatment 
of Kearfott types in the AMNH. 

satoi Kawabe, 1978 (Pseudohedya); Tinea 
10: 174. Holotype 6, Japan, Hokkaido, 
Mt. Tentosan, Abashiri, 4—5 August 
1975, R. Sato. 

sayonae Kawabe, 1985 (Acleris); Tinea 12: 
5. Holotype 6, Taiwan, Nantou Hsien, 


VOLUME 102, NUMBER 4 


Nanshanchi, 25—26 July 1983, A. Ka- 
wabe. 

saxicolana Walsingham 1879 (Cochlyis); 
Ill. Lepid. Heter. Brit. Mus. 4: 29. Syn- 
type 3d, USA, southern Oregon, end of 
May, 1872. Walsingham described this 
species from three males; the other two 
are presumed to be in the BMNH. 

scalaris Diakonoff, 1968 (Petrova); U.S. 
Natl. Mus. Bull 257: 86. Holotype &, 
Philippine Islands, Mountain Province, 
Luzon, base of Mt. Data, 5,500’, 22 
April 1946, CNHM Philippine Zool. 
Exped. 1946—47, H. Hoogstraal. 

scardiana Dognin, 1905 (Cnephasia); Ann. 
Soc. Entomol. Belgium 49: 86. Holo- 
type 2, Ecuador, El Monje de Loja, 
1894. 

schizodelta Diakonoff, 1982 (Grapholita); 
Zool. Verhandel. (Leiden) 193: 18. Ho- 
lotype ¢, Sri Lanka, Ratnapura District, 
Uggalkaltota, 350’, Irrigation Bunga- 
low, 31 January—8 February 1970, D. 
Davis & B. Rowe. 

schwarziana Busck, 1907 (Phalonia); J. N. 
Y. Entomol. Soc. 15: 25. Holotype °&, 
USA, Maryland, Montgomery Co., 
Plummers Island (in Potomac River 
above Washington, D.C.), July 1903, A. 
Busck. 

scleroductus Brown, 1991 (Punctapinella); 
Los Angeles Co. Mus. Contrib. Zool. 
423: 4. Holotype °, Brazil, Rio de Jan- 
iero. 3 July 1929, J. Zikan- 

sedatana Busck, 1906 (Hemimene); Proc. 
Biol. Soc. Wash. 19: 177. Holotype <6, 
USA, Colorado, South Park, 27 August 
1905. 

segregata Heinrich, 1924 (Thiodia); J. 
Wash. Acad. Sci. 14: 388. Holotype 6, 
USA, California, Tulare Co., Monachee 
Meadows, “July 16—23.” 

seiugata Razowski, 1987 (Terinebrica), 
Tinea 12(suppl.): 138. Holotype °, 
Peru, Cusco, Machu Picchu, 2,385 m, 4 
February 1959, J. Clarke. 

semicirculana Fernald, 1882 (Tortrix); 
Trans. Am. Entomol. Soc. 10: 68. Lec- 
totype 6, USA, Colorado, Morrison, 


1059 


1880. Designated by Brown and Powell 
@g91). 

senatrix Heinrich, 1924 (Exentera); J. 
Wash. Acad. Sci. 14: 390. Holotype ¢, 
USA, Arizona, Cochise Co., Paradise, 
*“March 8-15.” 

senilis Razowski, 1987 (Silenis); Tinea 
12(suppl.): 128. Holotype 6, Bolivia, 
Cochabamba, Incachaca, tropical cloud 
area, 17 August-5 September 1956, 
2,100 m, L. Pena. 

seorsa Heinrich, 1924 (Epinotia); J. Wash. 
Acad. Sci. 14: 392. Holotype 6, Cana- 
da, British Columbia, Vavenby, 16 Sep- 
tember 1922, T. Moilliet. 

separationis Heinrich, 1923 (Epiblema 
praesumptiosa); Bull. U.S. Natl. Mus. 
123: 143. Holotype ¢, USA, Texas, 
Cameron Co., Brownsville, 27 May 
1917, A. Busck. 

seraphicana Heinrich, 1923 (Hystricophora 
asphodelana); Bull. U.S. Natl. Mus. 
123: 258. Holotype 6, USA, Washing- 
ton, Whitman Co., Pullman, 13 May 
1898, C. Piper. Heinrich (1923) consis- 
tently misspelled Hystrichophora in his 
revision. 

serapicana Heinrich, 1923 (Eucosma); 
Bull. U.S. Natl. Mus. 123: 266. Holo- 
type 6, USA, Montana, Cascade Co., 
Great Falls, 8 July 1921, H. Dyar. 

serena Clarke, 1968 (Cochylis); Proc. U.S. 
Natl. Mus. 125: 20. Holotype 6, Brazil, 
Santa Catarina, Nova Teutonia, July 
1963, E Plaumann. 

sertula Diakonoff, 1982 (Licigena); Zool. 
Verhandel. (Leiden) 193: 16. Holotype 
6, Sri Lanka, Kandy, Udawattekelle, 
15-24 January 1970, D. Davis & B. 
Rowe. 

shastana Walsingham, 1879 (Paedisca); Il. 
Lepid. Heter. Brit. Mus. 4: 46. Syntype 
6, USA, California, Siskiyou Co., 
Mount Shasta, August 1871. This spe- 
cies was described from two males: one 
syntype in the USNM and the other ap- 
parently in the BMNH. 

shastensis McDunnough, 1955 (Anchylo- 
pera columbiana); Am. Mus. Novit. 


1060 


1725: 12. Holotype 6, USA, California, 
Siskiyou Co., Shasta Retreat, “‘June 8— 
Le 

shibatai Kawabe, 1985 (Archips); Tinea 12: 
1. Holotype 3, Taiwan, Chiai Hsien, 
Fenchihu, 1,400 m, 3 August 1973, Y. 
Shibata. 

shikokuensis Kawabe, 1984 (Epinotia); Tin- 
ea 11: 187. Holotype 6, Japan, Shiko- 
ku, Kagawa Pref., Oku-shio-iri, 5 Au- 
gust 1972, H. Toshima. 

shimekii Kawabe, 1974 (Zeiraphera); Tyo 
to Ga 25: 97. Holotype 6, Japan, Gun- 
ma Pref., Yunotaira, 26 July 1968, A. 
Kawabe. 

sicaria Diakonoff, 1982 (Acroclita); Zool. 
Verhandel. (Leiden) 193: 55. Holotype 
36, Sri Lanka, Galle District, Kanneliya, 
200’, 15—17 October 1976, G. Hevel, R. 
Dietz, R. Karunaratne & D. Balasoori- 
ya. 

sierrae Blanchard & Knudson, 1983 (Eu- 
cosma); Proc. Entomol. Soc. Wash. 85: 
850. Holotype ¢, USA, Texas, Culber- 
son Co., Sierra Diablo Wildlife Man- 
agement Area, 6,400’, 30 August 1970, 
J. Franclemont. 

sierrae McDunnough, 1955 (Anchylopera 
simuloides); Am. Mus. Novit. 1725: 14. 
Holotype 6, USA, California, Tulare 
Co., Mineral King, “‘June 24-30.” 

signiferana Heinrich, 1923 (Epinotia); Bull. 
U.S. Natl. Mus. 123: 232. Holotype 6, 
USA, California, San Diego Co., San 
Diego, 14 November 1911, W. Wright. 

silvertonana Obraztsov, 1962 (Anopina); 
Am. Mus. Novit. 2082: 19. Holotype ¢d, 
USA, Colorado, San Juan Co., Silver- 
ton, “July 24-31.” 

silvertoniensis Heinrich, 1923 (Epinotia); 
Bull. U.S. Natl. Mus. 123: 214. Holo- 
type 6, USA, Colorado, San Juan Co., 
Silverton, “July 16—23.” 

simplex Diakonoff, 1968 (Ebodina); Bull. 
U.S. Natl. Mus. 257: 35. Holotype <4, 
Philippine Islands, Luzon, Mt. Makil- 
ing, Baker. 

simplex McDunnough, 1925 (Eucosma); 
Canad. Entomol. 57:21. Holotype °&, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


USA, Iowa, Woodbury Co., Sioux City, 
7 May 1916. 

simpsoni Busck, 1903 (Carpocapsa pomo- 
nella); Proc. Entomol. Soc. Wash. 5: 
236. Holotype 6, USA, Idaho, Ada Co., 
Boise, 8 August 1902, C. Simpson. 

simulata Heinrich, 1928 (Rhyacionia),; 
Proc. Entomol. Soc. Wash. 30: 62. Ho- 
lotype °, Japan, Yokahoma, r.f. Pinus 
thunbergii, em: 5 July 1924, A. Kariya. 

sinistra Heinrich, 1926 (Bactra); Bull. U.S. 
Natl. Mus. 132: 87. Holotype 2, USA, 
Louisiana, Sabine River Ferry, opposite 
Orange, Cornell University Lot 542, 
Sub 20, 20 June 1917. 

siskiyouensis Heinrich, 1923 (Epinotia pul- 
satillana); Bull. U.S. Natl. Mus. 123: 
202. Holotype 6, USA, California, Sis- 
kiyou Co., Shasta Retreat, “Aug 16— 


skinnerana Heinrich, 1923 (Suleima); Bull. 
U.S. Natl. Mus. 123: 157. Holotype 6, 
USA, Arizona, Cochise Co., Huachuca 
Mountain, Carr Canyon, August 1905, 
H. Skinner. 

sonae Clarke, 1949 (Eulia); Acta Zool. Lil- 
loana (Tucuman) 7: 587. Holotype ¢, 
Brazil, Santa Catarina, 12 July 1935, E 
Hoffmann. 

sotipena Brown, 1986 (Epinotia); J. Lepid. 
Soc. 40: 341. Holotype 2, USA, Mary- 
land, Montgomery Co., Plummers Is- 
land, Potomac River, 7 April 1962, R. 
W. Hodges. 

spaldingiana Obraztsov, 1961 (Argyrotaen- 
ia); Am. Mus. Novit. 2048: 20. Holo- 
type 6, USA, Utah, Utah Co., Provo, 4 
August 1908, T. Spalding. 

spartinana Barnes & McDunnough, 1916 
(Phalonia); Canad. Entomol. 48: 144. 
Holotype 36, USA, South Dakota, 
Union Co., Elk Point, rf. Spartina, 30 
July 1915, C. Ainslie. 

spectra Brown, 1992 (Catastega); J. N. Y. 
Entomol Soc. 100: 211. Holotype ¢, 
USA, Texas, Jeff Davis Co., Mt. Locke, 
McDonald Observatory, 19 July 1971, 
A. Blanchard. 

spinacea Razowski, 1994 (Psiathovalva); 


VOLUME 102, NUMBER 4 


SHILAP Revta. Lepid. 22: 69. Holotype 
3d, Venezuela, Aragua, Rancho Grande, 
1,100 m, 11—15 January 1966, S. & W. 
Duckworth. 

spinea Razowski, 1999 (Netechma); Polski 
Pismo Entomol. 68: 93. Holotype 6, 
Bolivia, Cochabamba, Incachaca, 2,100 
m, tropical cloud area, 27 August—5 
September 1956, L. Pena. Razowski 
(1999) incorrect identifies the disposi- 
tion of the type as NHMW. 

spiraeifoliana Heinrich, 1923 (Polychros- 
is); Proc. Entomol. Soc. Wash. 25: 106. 
Holotype 6, USA, Pennsylvania, Luz- 
erne Co., Hazelton, rf. Spiraea salici- 
yolia, -5/29-19e%, 

spirographa Diakonoff, 1968 (Bubonox- 
ena); Bull. U.S. Natl. Mus. 257: 66. Ho- 
lotype ¢, Philippine Islands, Luzon, Mt. 
Makiling. 

stadiana Barnes & Busck, 1920 (Peronea); 
Contrib. Nat. Hist. Lepid. North Am. 4: 
217. Holotype 6, Canada, Ontario, Ot- 
tawa, 18 September 1905, C. Young. 

stauroma Diakonoff, 1968 (Polemograp- 
tis); Bull. U.S. Natl. Mus. 257: 33. Ho- 
lotype 6, Philippine Islands, Negros 
Oriental, Canlaon, 915 m, light trap, 20 
December 1959, L. Quate. 

stellifera Kawabe, 1978 (Hiroshiinoueana); 
Tinea 10: 181. Holotype ¢, Japan, Yak- 
ushima Island, Kurio, 7 August 1972, T. 
Watanabe. 

stenampyx Diakonoff, 1982 (Ancylis); Zool. 
Verhandel. (Leiden) 193: 64. Holotype 
36, Sri Lanka, Kandy District, Udawat- 
tekelle, 3 November 1966. 

stenotes Clarke, 1976 (Alcina); Insects of 
Micronesia 9: 21. Holotype 3, Micro- 
nesia, Guam, Tenjo, rf. Styphelia, 16 
May 1936, R. Usinger. 

stevensi Powell, 1980 (Decodes); Pacific 
Insects 22: 90. Holotype ¢, USA, Col- 
orado, Larimer Co., Owl Canyon, 12 
km NW Fort Collins, June 1977, R. Ste- 
vens. 

straminoides Grote, 1873 (Conchylis); Bull. 
Buttalo Soc. Natl Sek (ll66 Type od; 
USA, New York, Buffalo. According to 


1061 


Fernald’s unpublished list, two Grote 
specimens of this species were received 
by the USNM; we found only one. The 
single Grote specimen cited above is 
clearly labeled as the type. 

strianus Fernald, 1905 (Archips); Canad. 
Entomol. 37: 399. ““Type’’ 6, Canada, 
Quebec, A. Hanham. Fernald mentions 
3 specimens in his description: a male 
from London, Ontario; a female from 
New Hampshire; and a male from Que- 
bec. He indicated that the former two 
were in his personal collection and the 
last was in the USNM. 

striatana Brown, 1999 (Dimorphopalpa); 
Pan-Pac. Entomol. 75: 88. Holotype ¢, 
Venezuela, Aragua, Rancho Grande, 
1,100 m, 21—25 January 1966, S. & W. 
Duckworth. 

striatanoides Brown, 1999 (Dimorphopal- 
pa); Pan-Pac. Entomol. 75: 88. Holo- 
type 6, Ecuador, Carci, Maldonado, 
2,200 m, 6—11 January 1963, V. Becker. 

strigatella Brown, 1992 (Catastega); J. N. 
Y. Entomol. Soc. 100: 216. Holotype d, 
USA, Texas, Brewster Co., Big Bend 
National Park, Chisos Basin, 29 July 
1982, E. Knudson. 

strigosa Heinrich, 1926 (Aphania); Bull. 
U.S. Natl. Mus. 132: 118. Holotype <4, 
USA, Alaska, Dawson, 16 June 1916, 
B. Clark. 

strigulana Brown & Obrazstov, 1991 (Dor- 
ithia); in Brown and Powell, Univ. Ca- 
lif. Publ. Entomol. 111: 46. Holotype &, 
Mexico, Veracruz, Las Vegas [ = Las 
Vigas], 8,000’, 1887, W. Schaus. 

stygiana Dyar, 1903 (Thiodia); Proc. Ento- 
mol. Soc. Wash. 5: 230. Five syntypes 
(3 66,2 22), USA, Colorado, Jeffer- 
son Co., Golden, May—June, Dyar & 
Caudell; syntype 6, USA, Arizona, 
Williams. This species was described 
from the six specimens cited above. 
One male from Colorado was placed in 
the type collection, presumably by J. 
Clarke; the other specimens are in the 
main collection. 

suadana Heinrich, 1923 (Eucosma); Bull. 


1062 


U.S. Natl. Mus. 123: 130. Holotype 6, 
USA, Utah, Vineyard, 6 August 1912, 
T. Spalding. 

subcandida Heinrich, 1929 (Thiodia for- 
mosana); Proco\U:S: Nath Mus...75;: 1. 
Holotype ¢, Canada, Alberta, Bilby, 12 
June 1924, O. Bryant. 

subditiva Heinrich, 1929 (Eucosma tahoen- 
sis): sProc: US: Nath Muss 75219» Ho- 
lotype d6, USA, California, Lake Tahoe, 
Deer Park Springs, “‘June 24-30.” 

subelectana Kawabe, 1976 (Olethreutes); 
Tinea 10: 46. Holotype ¢, Japan, 
Ohmaki, Toyama Pref., 3 June 1951, M. 
Hoshino. Although the specimen is la- 
beled “Olethreutes pseudoereciana,”’ it 
is clearly the specimen referred to in the 
original description. 

subflavana Walsingham, 1879 (Paedisca); 
Ill. Lepid. Heter. Brit. Mus. 4: 48. Syn- 
type d, USA, California. According to 
the original description, this species was 
described from two males and one fe- 
male from Rouge River, Oregon, end of 
May 1871. The USNM specimen bears 
a typical Walsingham type label and is 
presumed to be part of the original se- 
ries; we cannot explain the discrepancy 
in the locality data. 

subminimana Heinrich, 1923 (Thiodia); 
Bull. U.S. Natl. Mus. 123: 61. Holotype 
3, USA, California, San Diego Co., San 
Diego, ““Aug 1-7.” 

subolivata Clarke, 1955 (Orthocomtis); 
Trans. Royal Entomol. Soc. London 
107: 148. Holotype 6, Costa Rica, Car- 
tago Province, Tuis, 5,800’, 28 August 
1908, W. Schaus. 

subretracta Kawabe, 1976 (Olethreutes); 
Tinea 10: 45. Holotype 6, Japan, To- 
kyo, Mt. Takao, 11 June 1952, M. 
Hoshino & T. Haga. 

subsolana Miller, 1978 (Larisa); J. Lepid. 
Soc. 32: 258. Holotype 6, USA, Arkan- 
sas, Washington Co., Devil’s Den State 
Park, 26 June 1966, R. Hodges. 

subretiniana Obraztsov, 1962 (Choristo- 
neura); Am. Mus. Novit. 2101: 9. Ho- 
lotype 6, USA, California, Tulare Co., 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Monachee Meadows, 8,000’, “July 16— 
23.7 

substitutionis Heinrich, 1923 (Gypsonoma); 
Bull. U.S. Natl. Mus. 123: 163. Holo- 
type d, Canada, Manitoba, Aweme, 27 
July 1905, Criddle. 

subtropica Miller, 1960 (Rhyacionia); J. 
Lepid. Soc. 14: 231. Holotype 2, USA, 
Florida, Okaloosa Co., Valparaiso, rf. 
Pinus, em: 26 May 1927, E. Gemner. 

subviridis Heinrich, 1929 (Epinotia); Proc. 
U.S. Natl. Mus. 75: 15. Holotype <d, 
USA, California, San Diego Co., San 
Diego, 5 July 1921 [genitalia slide only; 
pinned specimen has not been found]. 
A paratype specimen is also in the type 
collection. 

sugii Kawabe, 1976 (Epiblema); Tinea 10: 
42. Holotype ¢6, Japan, Tokyo, Seta- 
gaya, 16 June 1963, A. Kawabe. 

sugii Kawabe, 1989 (Dichrorampha); Tinea 
12: 196. Holotype d, Taiwan, Hualien 
Hsien, Kuanyuan, 2,400 m, 7-8 August 
1987, A. Kawabe. 

sulawesiensis Kawabe, 1993 (Bathypluta); 
Tyo to Ga 43: 237. Holotype 2, N Su- 
lawesi, Tandano, December 1988. 

sulphurica Razowski, 1999 (Netechma); 
Polski Posmo Entomol. 68: 95. Holo- 
type 2, Brazil, Santa Catarina, 21 Jan- 
uary 1936, EK Hoffman. Razowski 
(1999) incorrect identifies the disposi- 
tion of the type as NHMW. 

superba Razowski, 1988 (Clarkenia); Acta 
Zool. Cracov. 31: 406. Holotype 3, Co- 
lombia, Cauca, Paramo de Parace, Lake 
San Rafael, 3,570 m, 27 January 1959, 
J. Clarke. 

symphyla Razowski, 1984 (Polyortha); 
Acta Zool. Cracov. 27: 222. Holotype 
3, Bolivia, Cochabamba, Incachaca, J. 
Steinbach. 

synchysis Pogue, 1986 (Apolychrosis); in 
Cibrian-Tovar et al., Cone and Seed In- 
sects of the Mexican Conifers: 27. Ho- 
lotype 6, Mexico, Tlaxcala, Municipio 
de Terrenate, Villareal, rf., Abies reli- 
giosa, em: 10 January 1985. 

synneurana Barnes & Busck, 1920 (Amor- 


VOLUME 102, NUMBER 4 


bia); Contrib. Nat. Hist. Lepid. North 
Am. 4: 211. Holotype 2, USA, Arizo- 
na, Redington. 


taedana Miller, 1978 (Petrova); Ann. En- 
tomol. Soc. Am. 71: 337. Holotype °, 
USA, Florida, Seminole Co., Geneva, 
21 May 1967, R. Leuschner. 

taetera Razowski, 1984 (Histurodes); Acta 
Zool. Cracov. 27: 214. Holotype 6, 
Guatemala, Cerro Zunil, 4,000—5,000’, 
Champion. 

tahoensis Heinrich, 1923 (Eucosma); Bull. 
U.S. Natl. Mus. 123: 112. Holotype <6, 
USA, California, Lake Tahoe, Deer 
Park Springs, “July 8—15.” 

taiwana Kawabe, 1986 (Zeiraphera); En- 
tomol. Pap. Pres. Kurosawa, Tokyo: 79. 
Holotype ¢3, Taiwan, Chiai Hsien, Al- 
ishan, 2,200 m, 12 August 1974, Y. 
Kishida. 

taiwana Kawabe, 1992 (Acleris); Tinea 13: 
174. Holotype 6, Taiwan, Nantou 
Hsien, Tayuling, forest, 2,500 m, 9-18 
June 1980, D. Davis. 

taiwana Kawabe, 1992 (Neopotamia cryp- 
tocosma); Japan. Hetero. J. 170: 359. 
Holotype 6, Taiwan, Hualien Hsien, 
Kuanyuan, 2,400 m, 7-8 August 1987, 
A. Kawabe. 

taiwanensis Kawabe, 1985 (Archips); Tinea 
12: 3. Holotype ¢, Taiwan, Nantou 
Hsien, Lushan Spa, 1,200 m, 27-29 
July 1983, A. Kawabe. 

taiwanica Razowski, 1977 (Aethes cni- 
cana); Tyo to Ga 28: 37. Holotype 6, 
Taiwan, Tattaka, 6 June 1943, S. Issiki. 

taleana Grote, 1878 (Grapholitha); Canad. 
Entomol. 10: 54. Holotype d (only a 
forewing), USA, Illinois, T. Bean. The 
original description appears to have 
been based on a single male specimen. 

tamaensis Kawabe, 1974 (Epinotia); Tyo to 
Ga 25: 96. Holotype 6, Japan, Tokyo, 
Tama Hill, 22 April 1958, A. Kawabe. 

tambomachaya Razowski, 1988 (Seticosta); 
Acta Zool. Cracov. 31: 401. Holotype 
2, Peru, Cusco, Tambomachya, 3,690 
m, 3 February 1959, J. Clarke. 


1063 


tantilla Heinrich, 1926 (Satronia); Bull. 
U.S. Natl. Mus. 132: 17. Holotype 6, 
USA, Florida, Alachua Co., Archer, 4 
May 1882. 

tayulingensis Kawabe, 1986 (Dichroram- 
pha); Entomol. Pap. Pres. Kurosawa, 
Tokyo: 83. Holotype ¢, Taiwan, Hu- 
alien Hsien, Tayuling, 2,560 m, 2—3 Au- 
gust 1983, A. Kawabe. 

taxifoliella Busck, 1914 (Evetria); Proc. 
Entomol. Soc. Wash. 16: 146. Holotype 
2, USA, Montana, Missoula Co., hill at 
canyon east of Missoula, rf. P. taxifolia, 
24 April 1913, J. Brunner. 

telesocia Razowski, 1994 (Cylichneulia); 
SHILAP Revta. Lepid. 22: 68. Holotype 
3, Venezuela, Aragua, Rancho Grande, 
1,100 m, 16—19 January 1966, S. & W. 
Duckworth. 

temerana Busck, 1907 (Phalonia); J. N. Y. 
Entomol. Soc. 15: 28. Holotype ¢d, 
USA, Pennsylvania, Allegheny Co., 
Oak Station, 21 May 1906, E Marloff. 

tenebrica Heinrich, 1929 (Anchylopera); 
Proc. U.S. Natl. Mus. 75: 17. Holotype 
6, Canada, Alberta, Bilby, 12 June 
1924, O. Bryant. 

tenebrica Razowski, 1987 (Terinebrica); 
Tinea 12(suppl:): 135. Holotype 6, 
Peru, Cusco, Machu Picchu, 2,385 m, 6 
February 1959, J. Clarke. 

terminana Busck, 1907 (Hysterosia); J. N. 
Y. Entomol. Soc. 15: 33. Holotype 6, 
USA, Pennsylvania, Allegheny Co., 
Pittsburgh, H. Engel. 

tetropsis Busck, 1913 (Eucosma); Insect. 
Inscit. Menst. 1: 91. Holotype 6, British 
Guiana, Kirby, rf. guava, H. Moore. 

teutoniana Brown, 1999 (Dimorphopalpa); 
Pan-Pac. Entomol. 75: 89. Holotype 6, 
Brazil, Santa Catarina, Nova Teutonia, 
27°11'S, 52°23'W, 300-500 m, Septem- 
ber 1963, EK Plaumann. 

texasana Blanchard & Knudson, 1984 (An- 
opina); Proc. Entomol. Soc. Wash. 86: 
449. Holotype 6, USA, Texas, Jeff Da- 
vis Co., Mt. Locke, 6,700’, 26 April 
1981, E. Knudson. 

thelmae Diakonoff, 1968 (Capua); Bull. 


1064 


U.S. Natl. Mus. 257: 23. Holotype ¢, 
Philippine Islands, Mindanao, Bukid- 
non, Mt. Katanglad, 1,480 m, 27-31 
October 1959, L. Quate & C. Yashi- 
moto. Although the holotype is labeled 
“Capua thalmae,”’ the original descrip- 
tion uses the correct spelling of “‘thel- 
mae.” 

theodora Clarke, 1976 (Herpystis); Insects 
of Micronesia 9: 76. Holotype 36, Mi- 
cronesia, Kusaie, Mutunlik, 22 m, 24 
January—30 April 1953, J. Clarke. 

theta Brown, 1991 (Punctapinella); Los 
Angeles Co. Mus. Contrib. Zool. 423: 
4. Holotype @, Venezuela, Aragua, 
Rancho Grande, 1,100 m, 15—21 August 
1967, R. Poole. 

thylacophora Diakonoff, 1968 (Peridaeda- 
la); Bull. U.S. Natl. Mus. 257: 76. Ho- 
lotype <6, Philippine Islands, Davao 
Province, Mindanao, E. slope Mt. Mc- 
Kinley, 17 September 1946, CNHM 
Philippine Zool. Exped., H. Hoogstraal. 

tocullionana Heinrich, 1920 (Eucosma); 
Proc. U.S. Natl. Mus. 57: 59. Holotype 
3, USA, Connecticut, Lyme, rf. cones 
of Picea, em: 16 May 1916, A. Cham- 
plain. 

tokui Kawabe, 1974 (Didrimys); Tyo to Ga 
25: 101. Holotype ¢, Japan, Yakushima 
Island, Aiko-dake, 30 March 1972, T. 
Watanabe. 

tokui Kawabe, 1974 (Eudemopsis); Kontyu 
42: 389. Holotype 6, Japan, Yakushima 
Island, Mt. Aiko-dake, 8 June 1972, T. 
Watanabe. 

tonsilis Razowski, 1984 (Ardeutica); Acta 
Zool. Cracov. 27: 217. Holotype 4, 
Costa Rica, Cartago Province, Juan Vi- 
nas. 

tonto Powell, 1980 (Decodes); Pacific In- 
sects 22: 93. Holotype 2, USA, Arizo- 
na, Pinal-Gila Co. line, Miami Superior 
Highway, 4,790’, 1-15 April 1925, O. 
Poling. 

toreuta Grote, 1873 (Penthina); Bull. Buf- 
falo Soc. Nat. Sci. 1: 92. Holotype ¢, 
USA, Pennsylvania. 

tornosema Clarke, 1968 (Phalonia); Proc. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


U.S. Natl. Mus. 125: 12. Holotype 9, 
Guatemala, Volcan Santa Maria, June, 
Schaus and Barnes. 

torostoma Clarke, 1972 (Laspeyresia); 
Proc. Entomol. Soc. Wash. 74: 467. Ho- 
lotype 2, Costa Rica, Cartago Province, 
Turrialba, r.f. string bean stems, 8 
March 1965, L. Bonnefil. 

toshimai Kawabe, 1974 (Eudemopsis); 
Kontya 42: 389. Holotype ¢, Japan, 
Kochi Pref., Mt. Ashizuriyama, Cape 
Ashizuri, 27 May 1973, H. Toshima. 

toshimai Kawabe, 1978 (Griselda); Tinea 
10: 186. Holotype 3, Japan, Shikoku, 
Kagawa Pref., 15 July 1972, T. Toshi- 
ma. 

towadaensis Kawabe, 1978 (Statherotis); 
Tinea 10: 176. Holotype 3, Japan, Hon- 
shu, Aomori Pref., Towada, 3 August 
1969, Sato & Oshima. 

trema Brown & Obraztsov, 1991 (Cuprox- 
ena); in Brown and Powell, Univ. Calif. 
Publ. Entomol. 111: 56. Holotype 6, 
Mexico, Guerrero, December 1920, R. 
Muller. 

triangulana Brown, 1992 (Catastega); J. N. 
Y. Entomol. Soc. 100: 220. Holotype 6, 
USA, Arizona, Gila Co., 4 mi ESE Pine, 
4,500 ft, 1 September 1961, R. W. 
Hodges. 

triangulana Kearfott, 1908 (Tortrix); J. N. 
Y. Entomol. Soc. 16: 179. Lectotype <6, 
USA, California, San Diego Co., San 
Diego, 31 July 1907. Designated by Ob- 
raztsov (1962). 

tricia Clarke, 1976 (Heleanna physalodes); 
Insects of Micronesia 9: 17. Holotype 
6, Micronesia, Kapingamarani Atoll, 
Hare Island, rf. Callophyllum inophyl- 
lum, 3 August 1946, Townes. 

tridens Razowski, 1988 (Seticosta); Acta 
Zool. Cracov. 31: 400. Holotype 3, Co- 
lombia, Cauca, Paramo de Parace, Lake 
San Rafael, 3,570 m, 27 January 1959, 
J. Clarke. 

trifida Adamski, 1986 (Apotomis); Canad. 
Entomol. 118: 666. Holotype ¢, USA, 
Washington, Ferry Co., Sherman Pass, 
23 July 1962, J. Clarke. 


VOLUME 102, NUMBER 4 


triopis Diakonoff, 1959 (Cryptaspasma); 
Zool. Verhandel. (Leiden) 43: 32. Ho- 
lotype ¢, Guam Island, Commander 
Mariana’s Hill, March 1949, K. Maeh- 
ler. 

triphera Brown & Obraztsov, 1991 (Cu- 
proxena); in Brown and Powell, Univ. 
Calif. Publ. Entomol. 111: 56. Holotype 
6, Panama, Cabima, May 1911, A. 
Busck. 

triquetra Obraztsov, 1964 (Proeulia); Proc. 
U.S. Natl. Mus. 116: 189. Holotype <d, 
Chile, Nuble, Chillan, 10 November 
1961, H. Lister. 

tristriata Kearfott, 1907 (Sparganothis); 
Trans. Am. Entomol. Soc. 33: 67. Syn- 
type ¢, USA, Minnesota, Duluth. Un- 
published lectotype selected by R. Lam- 
bert, labeled by J. Powell. Klots (1942) 
indicated that there was a female “‘co- 
type’’ in the AMNH, but did not des- 
ignate it as the lectotype. 

tsugae Issiki, 1961 (Epagoge); in Issiki and 
Mutuura, Microlepidoptera injurious to 
coniferous plants in Japan: 36. Holotype 
6, Japan, Osaka, Ikeda Pref., 23 May 
1958, T. Kodama. Komai (1999) unnec- 
essarily designated a lectotype for this 
species; apparently he was unaware that 
the holotype was deposited in the 
USNM. 

tsuifengana Kawabe, 1992 (Acleris); Tinea 
13: 175. Holotype 3, Taiwan, Nantou 
Hsien, Tsuifeng, 2,400 m, 29 December 
1989, A. Kawabe. 

tsushimaensis Kawabe, 1978 (Hedya); Tin- 
ea 10: 173. Holotype ¢, Japan, Tsush- 
ima Island, Nenbutsuzaka, 27 June 
1973, T. Watanabe. 

turbula Clarke, 1968 ([razona); Proc. U.S. 
Natl. Mus. 125: 40. Holotype °, Gua- 
temala, Volcdn Santa Maria, July, 
Schaus and Barnes. 

turialba Busck, 1920 (Hysterosia); Insect. 
Inscit. Menst. 8: 86. Holotype ¢, Costa 
Rica, Cartago Province, Juan Vinas, 
February, W. Schaus. 

twila Clarke, 1955 (Orthocomotis); Trans. 
Royal Entomol. Soc. London 107: 152. 


1065 


Holotype 6, Brazil, Santa Catarina, 
Nova Teutonia, February 1948, F Plau- 
mann. 


ulteriorana Heinrich, 1920 (Evetria); Proc. 
U.S. Natl. Mus. 57: 55. Holotype <4, 
USA, Oregon, Waldo, r.f. Psuedotsuga 
taxifolia, 16 August 1914, Keen & Ser- 
gent. 

umbrana Barnes & Busck, 1920 (Spargan- 
othis); Contrib. Nat. Hist. Lepid. North 
Am. 4: 212. Holotype 6, USA, Colo- 
rado, Adams Co., Denver, Oslar. 

umbraticana Heinrich, 1923 (Thiodia); 
Bull. U. S. Natl. Mus. 123: 70. Holo- 
type 6, USA, Colorado, Jefferson Co., 
foothills above Golden, 13 March 1901, 
Dyar & Caudell. 

unciana Dognin, 1904 (Atteria); Ann. Soc. 
Entomol. Belgium 48: 133. Holotype °, 
Ecuador, Zamora, September 1886. 

undata Walsingham, 1914 (Tortrix); Biol. 
Centr.-Am., Lepid., Heter. 4: 286. Ho- 
lotype 6, Mexico, Puebla, Popocatepetl 
Park, 8,000’, June 1906, W. Schaus. 

unguiculus Clarke, 1951 (Episimus); J. 
Wash. Acad. Sci. 41: 297. Holotype °, 
Argentina, Tigre, March 1946, E Bour- 
quin. 

unica Heinrich, 1923 (Epinotia); Bull. U.S. 
Natl. Mus. 123: 221. Holotype ¢, USA, 
Tennessee, Knox Co., Knoxville, rf. 
Bradburya virginiana, G. Ainslie. 

urbana Busck, 1912 (Tortrix); Proc. Ento- 
mol. Soc. Wash. 14: 86. Holotype <6, 
Mexico, Mexico City, November 1911, 
R. Muller. 

ustulatana Blanchard & Knudson, 1983 
(Eucosma); Proc. Entomol. Soc. Wash. 
85: 849. Holotype 6, USA, Texas, 
Washington Co., Brenham, 4 June 1979, 
E. Knudson. 

usuiana Kawabe, 1976 (Epibactra); Tinea 
10: 39. Holotype 6, Japan, Gunma 
Pref., Usui Pass, 5 June 1951, M. Hosh- 
ino. 

usurpata Razowski, 1987 (Utrivalva); Bull. 
Polish Acad. Sci. 35: 67. Holotype 6, 


1066 
Guatemala, Volcan Santa Maria, W. 
Schaus. 

uta Clarke, 1953 (Eucosma); J. Wash. 
Acad. Sci. 43: 226. Holotype 3, USA, 
Illinois, Putnam Co., 10 July 1938, M. 
Glenn. 


vaccinii Clarke, 1971 (Dichelopa); Smith- 
sonian Contrib. Zool. 56: 107. Holotype 
3, Rapa Island, Maugaoa, 950’ (292 
m), rf. Veronica rapensis, 5 November 
1963, J. & T. Clarke. 

vagana Heinrich, 1923 (Epinotia); Bull. 
U.S. Natl. Mus. 123: 230. Holotype ¢, 
USA, Washington, Liaga, rf. Pyrus ri- 
vularis [wild crab apple], 20—24 July 
1918, em: 20 July 1918, E. Heckard. 

variana Fernald, 1886 (Teras); Bull. U.S. 
Dept. Agric. Entomol. 12: 17. ““Type”’ 
3, USA, Maine. This is the only ex- 
ample of this species we could locate 
from the Fernald Collection. Fernald 
did not indicate how many specimens 
he examined. 

varnicosa Brown, 2000 (Lobogenesis); 
Proc. Entomol. Soc. Wash. 102: 36. Ho- 
lotype 2, Argentina, Tucuman, Ciudad 
Universitaria, 17 February 1959, J. 
Clarke. 

venatana Kawabe, 1992 (Acleris); Tinea 
13: 178. Holotype 6, Taiwan, Nantou 
Hsien, Tsuifeng, 2,400 m, 29 December 
1989, A. Kawabe. 

verecundana Blanchard, 1979 (Phaneta); J. 
Lepid. Soc. 33: 210. Holotype ¢, USA, 
Texas, Hemphill Co., Canadian, 15 Au- 
gust 1971, A. & M. Blanchard. 

versicolor Powell, 1978 (Rhyacionia); in 
Powell and Miller, U.S. Dept. Agric., 
Agric. Handb. 514: 30. Holotype dé, 
USA, Arizona, Coconino Co., Fort Val- 
ley, 7.5 mi NW Flagstaff, 29 June 1961, 
R. Hodges. 

vesta Clarke, 1968 (Phalonia); Proc. U.S. 
Natl. Mus. 125: 14. Holotype ¢, Ven- 
ezuela, Aragua, Rancho Grande, 1,100 
m, 16—23 October 1966, S. & W. Duck- 
worth. 

victoriana Busck, 1921 (Cacoecia); Canad. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Entomol. 53: 278. Holotype ¢, Canada, 
British Columbia, Victoria, 22 July 
1920, E. Blackmore. 

vietnamna Kawabe, 1993 (Cerace); Tyo to 
Ga 44: 99. Holotype 2, North Vietnam, 
Mt. Tam Dao, 20-31 July 1990, Mizun- 
uma. 

villana Busck, 1907 (Hysterosia); J. N. Y. 
Entomol. Soc. 15: 34. Holotype 6, 
USA, Colorado, Adams Co., Denver, 21 
June 1905, Oslar. 

violetana Kawabe, 1964 (Philedone); Tyo 
to Ga 15: 3. Holotype ¢, Japan, Hon- 
shu, Tokyo, Nippara, 16 September 
1963, A. Kawabe. 

virga Clarke, 1949 (Eulia); Acta Zool. Lil- 
loana (Tucuman) 7: 585. Holotype <6, 
Brazil, Santa Catarina, Nova Teutonia, 
2711'S, 52°23’ W, February 194835E 
Plaumann. 

virginiana Busck, 1914 (Evetria); Proc. En- 
tomol. Soc. Wash. 16: 145. Holotype @, 
USA, Virginia, Fairfax Co., Falls 
Church, rf. Pinus virginiana, em: 4 
May 1914, C. Heinrich. 

viridana Barnes & Busck, 1920 (Platyno- 
ta); Contrib. Nat. Hist. Lepid. North 
Am. 4: 213. Holotype 6, USA, Arizo- 
na, Yavapai Co., Dewey, ““May 1-7.” 

virifloscula Brown, 1991 (Cuproxena); in 
Brown and Powell, Univ. Calif. Publ. 
Entomol. 111: 67. Holotype 3, Vene- 
zuela, Aragua, Rancho Grande, 1,100 
m, 8-14 August 1967, R. Poole. 

vitrana Walsingham, 1879 (Grapholitha) 
Ill. Lepid. Heter. Brit. Mus. 4: 65. Syn- 
type 6, USA, Oregon (northern), April 
1872. According to the original descrip- 
tion, this species was described from 
four males; the other syntypes are pre- 
sumed to be in the BMNH. 


wellingtoniana Kearfott, 1907 (Olethreu- 
tes); Trans. Am. Entomol. Soc. 33:13. 
Lectotype 6, Canada, British Columbia, 
Wellington, 19 May, G. W. Taylor. Des- 
ignated by Blanchard & Knudson 
(1984). 

wielgusi Clarke, 1990 (Argyrotaenia); J. 


VOLUME 102, NUMBER 4 


Res. Lepid. 28: 97. Holotype 2, USA, 
Arizona, Cochise Co., Huachuca Mts., 
Pueblo del Sol, 6 November 1986, R. 
Wielgus. 

willettana J. Comstock, 1939 (Carolella 
busckana); Bull. South. Calif. Acad. 
Sein 353 115. Holotype *6 “USA, (Cali- 
fornia, Riverside Co., Chuckwalla 
Springs, 25 December 1937, G. Willett. 

worthingtoniana Fernald, 1878 (Paedisca); 
Canad. Entomol. 10: 83. Lectotype <4, 
USA, ‘North Illinois.”’ Designated by 
Miller (1970). 

wrighti Brown & Powell, 2000 (Anopina), 
Univ. Calif. Publ. Entomol. 120: 98. 
Holotype 6, USA, Colorado, Chaffee 
Co., 2.5 mi ESE Buena Vista, 23 Au- 
gust 1997, D. Wright. 


yama Kawabe, 1974 (Olethreutes bipunc- 
tana); Tyo to Ga 25: 102. Holotype 4, 
Japan, Hokkaido, Mt. Daisetsu, 23 July 
1959, M. Suzuki. 

yasudai Kawabe, 1972 (Cryptophlebia); 
Tinea 9: 242. Holotype 6, Japan, Sai- 
tama Pref., Owa, 7 August 1952, T. Ha- 
ruta. 

yasutoshii Kawabe, 1985 (Acleris); Tinea 
12: 9. Holotype 6, Taiwan, Taichung 
Hsien, Lishan, 24 August 1970, Y. Shi- 
bata. 

yuccatana Busck, 1907 (Phalonia); J. N. Y. 
Entomol. Soc. 15: 23. Holotype 2°, 
USA, Texas, Nueces Co., Nuecestown, 
Ei “Yucca baccata; W2»April 1897, C. 
Marlatt. 

yakushimensis Kawabe, 1974 (Notocelia), 
Tyo to Ga 25: 96. Holotype ¢, Japan, 
Yakushima Island, Onoaida, 7 October 
1972, T. Watanabe. 


zacualpana Busck, 1913 (Phtheochroa), In- 
sect. Incit. Menst. Wash. 1: 141. Holo- 
type 2, Mexico, Zacualpan, July 1913, 
R. Muller. 

zeteles Razowski, 1987 (Chicotortrix); Tin- 
ea 12(suppl.): 124. Holotype 6, Colom- 
bia, Bogota, Chico, 25 January 1959, J. 
Clarke. 

zimapanus Powell, 1980 (Decodes); Pacific 


1067 


Insects 22: 107. Holotype 2, Mexico, 
Hidalgo, 3 mi E Zimapan, 6,400’, 31 
July—1 August 1963, W. Duckworth & 
D. Davis. 

zona Clarke, 1986 (Dichelopa); Smithson- 
ian Contrib. Zool. 416: 144. Holotype 
36, Marquesas Islands, Nuku Hiva, Tap- 
uaooa, 2,500’, 30 January 1968, J. & T. 
Clarke. 


ACKNOWLEDGMENTS 


This list was begun by Brown in 1988 
during a one-year Smithsonian Post-doctor- 
al Fellowship, with the encouragement and 
assistance of the late J. KE Gates Clarke. 
From 1989 to about 1993 Clarke continued 
to supply the names of type specimens as 
they slowly surfaced from the USNM col- 
lection or were returned by colleagues. Jack 
had a strong ‘“‘collection-bias,’’ and we be- 
lieve he would have been pleased to see the 
USNM type collection thoroughly orga- 
nized and catalogued. 

We thank the following for helpful com- 
ments on the list: William Miller, Univer- 
sity of Minnesota, St. Paul; Eric Grissell 
and David Smith, Systematic Entomology 
Laboratory, USDA, National Museum of 
Natural History, Washington, DC; Richard 
Brown, Mississippi Entomological Muse- 
um, Mississippi State University, Mississip- 
pi; Kevin Tuck, The Natural History Mu- 
seum, London, England. 


LITERATURE CITED 


Blanchard, A. 1979. New status for Epiblema minu- 
tana (Kearfott) and new species of Epiblema Htib- 
ner and Sonia Heinrich (Tortricidae). Journal of 
the Lepidopterists’ Society 33: 179-188. 

Blanchard, A. and E. Knudson. 1984. Three new tor- 
tricids (Lepidoptera) from Texas. Proceedings of 
the Entomological Society of Washington 86: 
446-451. 

Brown, J. W. 1998. A new genus of tortricid moths 
from Chile related to Varifula Razowski (Lepi- 
doptera: Tortricidae). Journal of the Lepidopter- 
ists’ Society 52: 177-181. 

Brown, J. W. and J. A. Powell. 1991. Systematics of 
the Chrysoxena group of genera (Lepidoptera: 
Tortricidae: Euliini). University of California Pub- 
lications in Entomology 111: 1—87. 


1068 


Caudell, A. N. 1902. Some insects from the summit 
of Pike’s Peak, found on snow. Proceedings of the 
Entomological Society of Washington 5: 74-83. 

Clarke, J. E G. 1955. Neoptropical moths of the genus 
Orthocomotis Dognin (Lepidoptera: Tortricidae). 
Transactions of the Royal Entomological Society 
of London 107: 139-168. 

1968. Neotropical Microlepidoptera, XVII. 

Notes and new species of Phaloniidae. Proceed- 

ings of the United States National Museum 125: 

1-58. 

. 1971. The Lepidoptera of Rapa Island. Smith- 

sonian Contributions to Zoology 56: 1—282. 

. 1974. Presidential Address—1973. The Na- 

tional Collection of Lepidoptera. Journal of the 

Lepidopterists’ Society 28: 181—204. 

. 1976. Insects of Micronesia. Microlepidop- 
tera: Tortricoidea, Vol. 9(1). Bernice P. Bishop 
Museum. Honolulu, HI. 

Davis, D. 1985. Donation of Blanchard Lepidoptera 
Collection to the Smithsonian Institution. Journal 
of the Lepidopterists’ Society 39: 235-236. 

. 1996. Donation of the Atsushi Kawabe Lep- 
idoptera collection to the Smithsonian Institution. 
Journal of the Lepidopterists’ Society 50: 270— 
PD TPAN 

Diakonoff, A. 1968. Microlepidoptera of the Philip- 
pine Islands. Bulletin of the United States Nation- 
al Museum 257: 1—484. 

1982. On a collection of microlepidoptera 
from Sri Lanka (Ceylon). Zoologische Verhande- 
lingen (Leiden) 193: 1-124. 

Dyar, H. G. 1903 [1902]. A list of North American 
Lepidoptera. Bulletin of the United States Nation- 
al Museum 52: 1-723. 

Freeman, T. N. 1958. The Archipinae of North Amer- 
ica (Lepidoptera; Tortricdae). The Canadian En- 
tomologist 90(supplement): 1—89. 

Heinrich, C. 1923. Revision of the North American 
moths of the subfamily Eucosminae of the family 
Olethreutidae. Bulletin of the United States Na- 
tional Museum 123: 1-298. 

Heinrich, C. 1926. Revision of the North American 
moths of the subfamilies Laspeyresiinae and 
Olethreutinae. Bulletin of United States National 
Museum 132: 1—207. 

Klots, A. B. 1942. Type material of North American 
Microlepidoptera other than Aegeriidae in the 
AMNH. Bulletin of the American Museum of 
Natural History 79: 391—424. 

Komai, FE 1999. A taxonomic review of the genus Gra- 
pholita and allied genera (Lepidoptera: Tortrici- 
dae) in the Palaearctic region. Entomologica Scan- 
dinavica supplement 55, 226 pp. 

Miller, W. E. 1967. Taxonomic review of the Rhy- 
acionia frustrana group of pine-tip moths, with 
description of a new species (Olethreutidae). Ca- 
nadian Entomologist 99: 590-596. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


. 1970. Fernald types of North American Oleth- 

reutinae (Lepidoptera: Tortricidae). Proceedings of 

the Entomological Society of Washington 72: 

288-294. 

. 1973. Two previously unrecognized scientific 

names for the strawberry leafroller (Lepidoptera: 

Tortricidae). Annals of the Entomological Society 

of America 66: 553-554. 

. 1983. New synonymies in Nearctic Dichror- 
ampha (Lepidoptera: Tortricidae). Proceedings of 
the Entomological Society of Washington 85: 
727-733. 

Obraztsov, N. 1961. Descriptions of and notes on 
North and Central American species of Argyro- 
taenia, with the description of a new genus (Lep- 
idoptera, Tortricidae). American Museum Novi- 
tates 2048: 1—42. 

. 1962. Anopina, a new genus of the Cnephas- 

iini from the New World (Lepidoptera: Tortrici- 

dae). American Museum Novitates 2082: 1-39. 

. 1963. Some North American moths of the ge- 

nus Acleris (Lepidoptera: Tortricidae). Proceed- 

ings of the United States National Museum 114: 

213-270. 

. 1964. Neotropical Microlepidoptera, V. Syn- 

opsis of the species of the genus Proeulia from 

central Chile (Lepidoptera: Tortricidae). Proceed- 

ings of the United States National Museum 116: 

183-196. 

. 1965. On Paedisca radicana and the type spe- 

cies of the genus Griselda (Lepidoptera, Tortrici- 

dae, Olethreutinae). American Museum Novitates 

2213: 1-16. 

1966a. Neotropical Microlepidoptera, VIII. 

New genus Pseudomeritastis and its species (Lep- 

idoptera: Tortricidae). Proceedings of the United 

States National Museum 118: 221-232. 

. 1966b. Neotropical Microlepidoptera, IX. Re- 

vision of genus Pseudatteria (Lepidoptera: Tortri- 

cidae). Proceedings of the United States National 

Museum 118: 577-622. 

. 1966c. Neotropical Microlepidoptera, XI. Re- 
vision of genus /dolaterria (Lepidoptera: Tortri- 
cidae). Proceedings of the United States National 
Museum 119: 1-12. 

Razowski, J. 1999. Synopsis of Jcteralaria and Ne- 
techma Razowski (Lepidoptera: Tortricidae) with 
description of new taxa. Polskie Pismo Entomo- 
logiczne 68: 91—106. 

Riley, C. V. 1881. Descriptions of some new Tortrici- 
dae (leaf-rollers). Transactions of the St. Louis 
Academy of Science 4: 1-9. 

Rubinoff, D. and J. A. Powell. 1999. Description of 
Diedra, new genus (Lepidoptera: Tortricinae: Ar- 
chipini), and three new species, based on phylo- 
genetic analysis. Annals of the Entomological So- 
ciety of America 92: 473—487. 


VOLUME 102, NUMBER 4 


Smith, J. B. 1891. List of the Lepidoptera of Boreal 
North America. American Entomological Society, 
Philadelphia, 124 pp. 

Walsingham, Lord T. de Grey. 1979. Illustrations of 
typical specimens of Lepidoptera Heterocera in 
the collection of the British Museum. Part IV. 


1069 


North American Tortricidae. Trustees of the Brit- 
ish Museum, London. 

. 1914. Lepidoptera-Heteroptera, Vol. 4. Tinei- 
na, Pterophorina, Ornueodina, and Pyralidina and 
Hepialidina (part). Jn Biologia Centrali-America- 
na. Insects. British Museum, London. 482 pages. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 1070-1071 


NOTE 


Fitchia aptera Stal (Hemiptera: Reduviidae): 
Seasonal History and Habits in Mid-Appalachian Shale Barrens 


Fitchia aptera Stal is an infrequently col- 
lected though widely distributed harpactorine 
reduviid. Adults are 9-14 mm long, elongate, 
and dull yellow, with a dark longitudinal 
stripe on either side of the abdominal mid- 
line; this dorsal stripe is conspicuous in the 
brachypterous morph. Slater and Baranowski 
(1978. How to Know the True Bugs, W. C. 
Brown, Dubuque, Iowa. 256 pp.) illustrated 
the brachypterous and macropterous forms. 
Fitchia aptera can be confused with the only 
other member of this Nearctic genus, F. spi- 
nosula Stal; characters traditionally used to 
distinguish the two sympatric species—in- 
cluding presence or absence of prothoracic 
spines—are unreliable (McPherson et al. 
1992. Florida Entomologist 75: 222-230). 
McPherson et al. (1992) redescribed the 
adults and provided additional characters to 
separate these species. 

DeCoursey (1963. Bulletin of the Brook- 
lyn Entomological Society 58: 151—156) 
described and illustrated the immature stag- 
es and determined their duration by rearing 
F. aptera in the laboratory (“‘room temper- 
ature,’’ RH not stated); nymphs were fed 
small flies and homopterans. DeCoursey 
(1963) reported mean development (egg to 
adult) of 60 days (range 51—67; n not stat- 
ed) and found that F. aptera has only four 
nymphal instars. Swadener and Yonke 
(1973. Journal of the Kansas Entomological 
Society 46: 123-136) reared this reduviid 
in the laboratory (21°C night, 29°C day; 
40—60% RH), reporting that total develop- 
ment averaged 70 days (range 60.5—74.5; n 
= 134 eggs, n = 3 instar IV). They also 
confirmed the presence of only four instars. 

In contrast to the availability of biological 
data from laboratory studies, little is known 
about the habits of F. aptera in nature. Rea- 
dio (1927. University of Kansas Science 


Bulletin 17: 5—291) did not treat the bionom- 
ics of this species in his studies on reduviid 
biology. This mainly ground-dwelling assas- 
sin bug occurs by day at the bases of grass 
clumps in old fields, the adults overwinter- 
ing under boards, stones, or at roots of grass- 
es. Adults occasionally can be swept from 
tall grasses in early evening. Nymphs are 
found on the ground, mainly in grassy areas 
(DeCoursey 1963). Hagerty and McPherson 
(2000. Great Lakes Entomologist 32: 133-— 
160) reviewed additional collection records 
and in Illinois reported an adult at the roots 
of a grass clump, three adults from sweeping 
herbaceous vegetation, and another adult 
from sweeping a grassy field. 

Here I give notes on the seasonality of 
F. aptera in shale barrens and outcrops in 
the mid-Appalachians. Collections and ob- 
servations were made while inventorying 
the plant bug (Hemiptera: Miridae) fauna of 
moss phlox (Phlox subulata L.) during 
1989-1995. Mats of this prostrate perennial 
(family Polemoniaceae) were shaken over a 
white tray (Wheeler. 1995a. Proceedings of 
the Entomological Society of Washington 
97: 435—451), and adults and all instars of 
F. aptera were collected; nymphs were de- 
termined to instar in the laboratory, using 
DeCoursey’s (1963) descriptions. Voucher 
specimens, including nymphs, are deposited 
in the National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 

Unless otherwise stated, F. aptera was 
collected from mats of moss phlox at the 
following localities. The numbers of 
nymphs (Roman numerals = instars) and 
adults (A) are given parenthetically after 
other collection data. 

Maryland: Allegany Co., Country Club 
shale barren, 2.5 km NE of Cumberland, 2 
July 1994 (1 IID). Pennsylvania: Bedford 


VOLUME 102, NUMBER 4 


Co., shale slope, Rt 56 E of Fishertown, 24 
June 1990 (1 I); shale slope, Rt. 96 NE of 
Ryot, 24 June 1990 (2 I) & 10 May 1992 
(1 A). Virginia: Bath Co., Fort Lewis shale 
barren, Rt. 678 at Cowpasture River, 26 June 
1994 (1 I); Highland Co., Head Waters shale 
barren nr. jct. Rts. 250 & 616, 23 June 1990 
(7 I-III), 25 Aug. 1990 (1 IV; 2 A ex undet. 
sprawling plant), 10 July 1993 (3 II; 2 II ex 
base of Eriogonum allenii S. Wats), 1 Aug. 
1935021 tt), 26 June 1994 1.1.1, 
6 Aug. 1994 (1 I, 1 ID, 26 Aug. 1994 (1 
IV); Rockbridge Co., Furnace Mountain 
shale barren, ca. 3 km S of Natural Bridge 
Station, 24 May 1995 (1 I, 1A ex Antennaria 
plantaginifolia (L.) Richardson); Rocking- 
ham Co., shale slope, For. Rd. 87, W of 
Fulks Run, 19 May 1994 (1 A; 1 A at base 
of Tephrosia virginiana (L.) Pers.), 25 June 
1994 (1 I); Shenandoah Co., Short Mountain 
shale barren, ca. 4.6 km SE of Mount Jack- 
son, 10 July 1993 (1 III). West Virginia: 
Greenbrier Co., shale slope, Whites Draft 
Rd., Alvon, 4 July 1994 (1 II); Hampshire 
Co., shale bank, Rt. 29, 5.6 km N of Sla- 
nesville, 30 June 1990 (1 II); Hardy Co., 
shale bank, Mathias, 30 June 1990 (1 ID). 

Insect-plant associations in shale barrens 
have been little studied (e.g., Wheeler 
1995a; Braunschweig et al. 1999. pp. 83- 
98. In Anderson, R. C. et al., eds., Savan- 
nas, Barrens, and Rock Outcrop Commu- 
nities of North America. Cambridge Uni- 
versity Press, UK), but these xeric com- 
munities of high soil-surface temperatures 
(sometimes 50—60°C; Braunschweig et al. 
1999) harbor great insect richness (Wheeler 
1995a, b. Virginia Journal of Science 46: 
148). The predacious F. aptera can be add- 
ed to the diverse insect fauna of moss phlox 
in mid-Appalachian shale barrens and out- 
crops (Wheeler 1995a, b). All collections 
were from the Valley and Ridge physio- 
graphic province except the one in Rock- 
bridge Co., Va., which was from the Blue 
Ridge province. According to McPherson et 
al. (1992), Maryland, Virginia, and West 
Virginia are new state records. 

Only small numbers of F. aptera were 


1071 


found during any 45—60 minutes’ sampling 
of moss phlox, the seven early instars on 23 
June 1990 representing the most individuals 
in any sample. Adults, all brachypters (n = 
6), were found at four sites where nymphs 
were present; nymphs at other sites proba- 
bly also were F. aptera rather than F. spi- 
nosula. 

Overwintered adults were found under 
mats of moss phlox, at the base of Tephro- 
sia virginiana, or among stolons of Anten- 
naria plantaginifolia during May in shale 
barrens and outcrops. Except for the late- 
May collection in Rockbridge Co., Va., first 
instars were not observed before late June 
despite intensive sampling of moss phlox 
during April, May, and early June (Wheeler 
1995a). An early instar was observed feed- 
ing on an immature isopod at the Head Wa- 
ters shale barren in Virginia in late June. 
Third and fourth instars were present from 
early July to late August, suggesting a rath- 
er late-season, univoltine life cycle. Two 
adults found in late August likely were 
those of a new, rather than overwintered, 
generation. No nymphs or adults were ob- 
served during sampling of moss phlox in 
September and early October. 

In Connecticut, DeCoursey (1963) re- 
ported that overwintered adults of F. aptera 
become active in April. An adult collected 
in early April in Missouri did not oviposit 
in the laboratory until mid-May and contin- 
ued to lay eggs until mid-July (Swadener 
and Yonke 1973). Such observations, cou- 
pled with mean development of 60 (De- 
Coursey 1963) or 70 days (Swadener and 
Yonke 1973), also suggest that F. aptera is 
a relatively late-season reduviid. 

I thank Peter Adler (Clemson University) 
for comments that improved the manu- 
script, and Aaron Hagerty (Clemson Uni- 
versity) and Jay McPherson (Southern Illi- 
nois University) for allowing me to refer to 
information in a paper in press. 


A. G. Wheeler, Jr., Department of Ento- 
mology, Clemson University, Clemson, SC 
29634, U.S.A. (e-mail: awhlr@clemson. 
edu) 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 1072-1073 


NOTE 


First Host Plant and Parasitoid Record for Anastrepha spatulata Stone 
(Diptera: Tephritidae) 


Anastrepha spatulata Stone has been re- 
ported in the U.S. (S. Texas), Mexico, Cen- 
tral America (Guatemala S. to Panama), 
Venezuela and Tobago (Norrbom et al. 
1999. Myia 9: 65-251 [p. 82 for A. spatu- 
lata]). In Mexico, it is commonly captured 
in McPhail traps placed in commercial fruit 
orchards (Aluja et al. 2000. Proceedings of 
the Entomological Society of Washington 
102: 802-815). In the past 60 years, there 
have been considerable, but until now un- 
successful, efforts to discover its host plant. 
This species, together with A. alveata 
Stone, A. alveatoides Blanchard, A. distans 
Hendel, A. interrupta Stone, A. manihoti 
Lima, A. montei Lima, A. pickeli Lima, and 
A. umbrosa Blanchard, form the spatulata 
species group (Norrbom et al. 2000. pp. 
343-362. In Aluja and Norrbom, eds., Fruit 
Flies (Diptera: Tephritidae): Phylogeny and 
Evolution of Behavior, CRC Press). 

We report here on A. spatulata infesta- 
tions in Schoepfia schreberi Gmelin fruit 
(Olacaceae). This bushy plant is locally 
known in Veracruz as “guayabillo” or 
‘“‘palo meco.”’ Fully ripe fruits measure, on 
the average, 1.92 + 0.06 cm (diameter, n = 
25) and weigh 185 + 0.061 mg (n = 25). 
A mature plant measures ca. 2.10 m in 
height. Floral buttons of S. schreberi were 
observed from January 20th to February 
Sth. The flowering period was recorded 
from the Ist to the 12th of February and 
the fruiting season occurred between the 
10th and the 20th of February (all during 
1998). Fly larvae were always found feed- 
ing in the seed of the fruit (there is one seed 
per fruit). There was always only one larva 
per fruit. Given the small size of the host, 
fully mature larvae, 8.6 + 0.12 mm (n = 
8) in length, looked very compressed inside 
the host. We collected 3 batches (different 


collection dates) of ripe fruit from the 
ground adjacent to four S. schreberi bushes. 
A total of 583 g of fruit were collected on 
February 2, 9, and 18, 1998. Fruit was han- 
dled as described by Aluja et al. (2000). 
Pupae were harvested between 1 and 6 days 
after the fruit collecting date. The 583 g of 
fruit yielded 579 pupae (0.993 pupae/g of 
fruit). Mean pupal weight was 12 + 0.001 
mg (n = 13). We divided pupae into two 
batches. One was kept under field condi- 
tions and another was transported to the 
laboratory. The collection site was Llano 
Grande, Municipio de Teocelo, Veracruz, 
Mexico, at 720 m altitude (19°22’N and 
26°53'W). Native vegetation around this 
site is a tropical deciduous and subdeci- 
duous forest (Castillo-Campos. 1995. Ecol- 
ogia del Paisaje del Municipio de Jalco- 
mulco, Veracruz, MS Thesis, UNAM, Mex- 
ico City). 

Between February 29 and March 5, 454 
A. spatulata adults (223 females, 231 
males), 17 Doryctobracon areolatus (Sze- 
pligeti) (10 females, 7 males) and 2 Doryc- 
tobracon toxotrypanae (Muesebeck) (2 fe- 
males) emerged. We note that the D. areo- 
latus individuals emerging from A. spatu- 
lata were uniformly more darkly colored 
than those individuals of the same species, 
parasitizing, in the same region, A. alveata, 
A. obliqua (Macquart), A. striata Schiner 
and A. fraterculus (Wiedemann) (Lopez et 
al. 1999. Biological Control 15: 119-129). 
Length of the pupal period for flies was 22 
+ 3 and 27 + 9 days under laboratory and 
field conditions, respectively (mean tem- 
perature of 26 + 1°C and 23 + 3°C under 
laboratory and field conditions, respective- 
ly). In the case of the parasitoids, the length 
of the pupal period for D. areolatus was 25 
+ 9 and 26 + 5 days under laboratory and 


VOLUME 102, NUMBER 4 


field conditions, respectively (same temper- 
ature conditions as flies). For D. toxotry- 
panae it was 21 days (laboratory conditions 
only). Another 9 D. areolatus (8 females, 1 
male) entered diapause and emerged after 
ca. 11 months (range of 308-341 days). 
Diapause by D. areolatus in the same re- 
gion has been recently reported by Aluja et 
al. (1998. Annals of the Entomological So- 
ciety of America 91: 821-833). 

Schoepfia schreberi had been previously 
reported as a host plant of A. interrupta 
(McClanahan. 1951. State Plant Board of 
Florida Biennial Report No. 18, p. 44; as S. 
chrysophylloides). Interestingly, the host 
plant of another fly belonging to the same 
species group (A. alveata), also belongs to 
the family Olacaceae (Piedra et al. 1993. 
Proceedings of the Entomological Society 
of Washington 95: 127). Based on the few 
host records for the spatulata group (Norr- 
bom. 2000. Diptera Data Dissemination 
Disk 2, in press), it appears that flies in this 
assemblage of species are specialized on 
only two plant families, Olacaceae and Eu- 
phorbiaceae. 

Our discovery of D. toxotrypanae para- 
sitizing a species of Anastrepha is note- 
worthy since this parasitoid had been pre- 
viously reported parasitizing only T. curvi- 
cauda Gerstaecker (Wharton and Marsh. 
1978. Journal of the Washington Academy 
of Sciences 68: 147-167). Toxotrypana 
curvicauda is common in the study region 
where it has been recently reported infest- 
ing Gonolobus niger (Cav.) R. Br. (Ascle- 
piadaceae) fruit (Aluja et al. 2000). It is 
thus likely that the D. toxotrypanae individ- 
uals parasitizing A. spatulata stemmed from 
parasitized T. curvicauda. We note, how- 
ever, that D. toxotrypanae is only distin- 
guished from D. crawfordi on the basis of 
color differences (D. toxotrypanae is dark- 
er) (R. A. Wharton, pers. comm.). Based on 


1073 


the latter, the question remains open as to 
which of the two above mentioned Doryc- 
tobracon species was actually parasitizing 
A. spatulata in our study region. 

Voucher specimens of S. schreberi and A. 
spatulata are placed in the herbarium and 
insect collection of the Instituto de Ecolo- 
gia, A.C. in Xalapa, Veracruz, Mexico. An- 
astrephe spatulata individuals were also 
sent to the National Museum of Natural 
History, Washington, DC (A. L. Norrbom) 
and D. areolatus and D. toxotrypanae in- 
dividuals were sent to the parasitoid collec- 
tion of Texas A&M University, College 
Station, TX (R. A. Wharton). 

Specimen identification was by the fol- 
lowing: A. spatulata (A. L. Norrbom, Sys- 
tematic Entomology Laboratory, ARS, 
USDA, % National Museum of Natural 
History, Washington, DC); D. areolatus 
and D. toxotrypanae (Robert Wharton, Tex- 
as A & M University, College Station), and 
Schoepfia schreberi (Carlos M. Duran-Es- 
pinoza, Instituto de Ecologia, A.C., Xalapa, 
Veracruz, Mexico). Financial support was 
provided by the Campania Nacional contra 
las Moscas de la Fruta, Comisién Nacional 
para el Conocimiento y Uso de la Biodiv- 
ersidad (CONABIO) (Grant No. H296), 
Sistema Regional de Ciencia del Golfo 
(CONACyT-SIGOLFO) (Grant No. 96-01- 
003-V) and the U.S. Department of Agri- 
culture (ARS, USDA) (Grant Numbers 
198-23 and 58-6615-3-025). 


Martin Aluja, Emanuel Herrera, Maurilio 
Lépez, and John Sivinski (MA, EH, ML) 
Instituto de Ecologia, A.C., Apartado Post- 
al 63, 91000 Xalapa, Veracruz, Mexico (e- 
mail: alujam @ ecologia.edu.mx); (JS) 
Center for Medical, Agricultural, and Vet- 
erinary Entomology, Agricultural Research 
Service, U.S. Department of Agriculture, 
P.O. Box 14565, Gainesville, FL 32604, 
U.S.A. 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, p. 1074 


BoOoK REVIEW 


Aphids on the World’s Crops, An Iden- 
tification and Information Guide, Second 
Edition, by R. L. Blackman and V. EF Ea- 
stop. John Wiley & Sons, Baffins Lane, 
Chichester, Sussex PO19 1UD, England. 
466 pp. 2000. $145.00. 


The first edition of Aphids on the 
World’s Crops was published in 1984 and 
brought together information available pre- 
viously only in widely-distributed papers 
and publications containing species’ de- 
scriptions and keys. The information in the 
keys, in the systematic reviews, and in the 
photographs, will allow a non-specialist to 
identify, with a good degree of accuracy, 
aphids colonizing (feeding and producing 
young) the crops treated. 

The second edition has seven sections. 
Section A (pages 1—24) contains general in- 
formation on systematics, life cycles, host- 
plant relationships, geographical distribu- 
tion, and morphology and key characters. 
Section B (pages 25-214) begins with a 
statement on the limitations and use of the 
keys and is devoted to crops and their 
aphids. This section includes an alphabeti- 
cal listing of the Latin names of crops with 
their common names followed by an alpha- 
betical listing of the crops by common 
names, a list of the aphids reported to col- 
onize the crop, and keys to those aphids. 
Section C (pages 215-362) covers a dis- 
cussion of the aphids and includes a brief 
introduction followed by an alphabetical 
listing by genus and species of the aphids 
treated. This section is a wealth of infor- 
mation on aphids and includes field char- 
acters, host plants, virus transmission, dis- 
tribution, and biology. Section D (pages 
363-365) covers techniques on collecting, 
preserving and mounting, and labeling and 
storage. Section E (pages 367-373) con- 
tains sources of information on regionally 
classified faunal publications; general biol- 


ogy; morphology, anatomy, and physiolo- 
gy; genetics and development; migration 
and dispersal; relationships with other in- 
sects; host-plant relationships; control; and 
bibliographies. Section F (pages 375—414) 
is a list of references. Section G (pages 
415—466) is a photographic guide to slide- 
mounted aphids. 

While both editions are 466 pages in 
length, the second edition is about % larger 
in size, covers 38 more species, and con- 
tains more information in the keys and the 
systematic section. While the basic format 
is unchanged, the arrangement of the text 
and the collection of the illustrations in 
plates rather than their being shown with 
each key couplet is a more efficient use of 
space, is more pleasing to the eye, and does 
not detract from the value of or ease in fol- 
lowing the information. The biggest im- 
provement, from a user’s standpoint, is that 
the type in the section with the keys is larg- 
er and more legible. The photomicrographs 
and illustrations are much improved with 
the target characters easier to see. Having 
the names of the species with each photo- 
graphic plate is useful, but I missed having 
the listing of all species at the beginning of 
the photographic guide. 

Aphids on the World’s Crops remains the 
only current, world-wide reference with 
general and specific information on aphids 
colonizing crops and is a must have for 
anyone working in this field. Everyone who 
has a copy of the first edition will want to 
keep it as a reference (mine is well worn) 
but will want to purchase a copy of the sec- 
ond edition as a valuable working tool. 


Manya B. Stoetzel, Systematic Entomol- 
ogy Laboratory, PSI, Agricultural Research 
Service, U.S. Department of Agriculture, 
Bldg. 005, BARC-West, 10300 Baltimore 
Avenue, Beltsville, MD 20705, U.S.A. (e- 
mail: mstoetze @ sel.barc.usda.gov) 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 1075-1076 


Book REVIEW 


Classification of Lepidoptera. Part 1. In- 
troduction, by John B. Heppner. Holarc- 
tic Lepidoptera, Volume 5, Supplement 1. 
Published by the Association for Tropical 
Lepidoptera, Gainesville, FL. 1998. 148 
pp. + 6 pp. index. Soft bound, 8.5 X 11 
in. ISBN 1070-4140. $25.00 


This volume, the first of a proposed 
three-part ‘“‘Manual of Lepidoptera,” is 
chuck full of interesting factoids, keys, il- 
lustrations, and references. It presents a 
“new” higher classification of the order 
with novel (and sometimes unusual) group- 
ings of families based on the author’s ex- 
pertise and biases. Included are a tabular 
summary of Heppner’s (1991) previous 
work on species richness and biogeographic 
distribution of each of the superfamilies; 
‘schematic phylogenies” of most of the 
major groups (apparently based on intuition 
rather than analysis); an impressive data 
matrix (Table 2) of 24 morphological char- 
acters of adults, larvae, and pupae across all 
the superfamilies; an extensive key to 
adults of the major families, previously 
published by Heppner (1996); 33 pages of 
illustrations of wing venation of all major 
groups (never before compiled and pre- 
sented in such an easily browsed manner); 
and references, lots of them, subdivided by 
category (e.g., taxonomy, morphology, bi- 
ology, biogeography). The volume brings 
together widely scattered concepts and lit- 
erature, synthesized in Heppner’s unmistak- 
able style. 

If this book had been published 10 years 
ago, it might have stood for a decade as the 
standard reference on the order. Unfortu- 
nately, any impact that it potentially may 
have had likely will be overshadowed by 
the recently published Handbook of Zool- 
ogy (Kristensen 1999). Heppner’s timing 
could not have been worse. In contrast to 
Heppner’s single-authored manual, the edi- 
tor of the Handbook enlisted the assistance 


of 29 leading authorities in particular Lep- 
idoptera families, creating a work of grand 
proportions. 

Aside from being scooped by the Hand- 
book, Heppner’s work is plagued by a few 
quirky shortcomings and/or inconsistencies. 
Based on my knowledge of Tortricoidea (I 
have limited knowledge of the higher clas- 
sification of the entire order), I perused the 
volume for details of this superfamily. Ta- 
ble 1 indicates that it includes 6,683 spe- 
cies, with 250 species occurring in the Ethi- 
opian (Afrotropical) region. Actually, there 
are over 8,000 described species of Tortri- 
cidae (based on a checklist I have com- 
piled), and Diakonoff, in various publica- 
tions (e.g., Diakonoff 1960, 1970, 1973, 
1981, 1983), treated over 250 species of 
tortricids from Madagascar alone. Also, of 
ten genera of Tortricidae for which wing 
venation is illustrated (page 78), Heppner 
uses older, antiquated names for two: *“‘Tor- 
trix’ for Xenotemna (X. pallorana) and 
‘““Clysiana” for Eupoecilia (E. sp.). Given 
these minor errors in the group with which 
I am most familiar, I am skeptical about the 
quality of information presented for other 
superfamilies. 

In some cases, Heppner appears to have 
disregarded the published findings of other 
specialists. For example, Epstein’s (1996) 
analysis of the limacodid-group of families 
provides compelling evidence that Megal- 
opygidae, Aididae, Dalceridae, and Lima- 
codidae form a monophyletic group within 
Zygaenoidea. Yet in Heppner’s classifica- 
tion, Megalopygidae (including Aidinae) is 
placed in Zygaenoidea, while Dalceridae 
and Limacodidae are included in Cosso- 
idea. 

Despite its incongruities and shortcom- 
ings, Heppner’s compendium is a valuable 
source of information on the classification 
of Lepidoptera, and comparisons with the 
Handbook of Zoology are premature since 


1076 


parts 2 and 3 of Heppner’s work presum- 
ably will present morphological and biolog- 
ical details on each of the superfamilies, 
one of the strengths of the Handbook. Com- 
parisons with the Handbook also are unfair 
given the considerable difference in price— 
Heppner’s part 1 sells for about $25.00; the 
Handbook goes for a whopping $250! Be- 
cause currently proposed classifications of 
the Lepidoptera exhibit little consensus, all 
must be viewed as viable and reasonable 
alternative interim solutions. Heppner’s 
contribution will not go unnoticed. 


LITERATURE CITED 


Diakonoff, A. 1960. Tortricidae from Madagascar. Part 
I. Tortricinae and Chlidanotinae. Verhandelingen 
der Koninkluke Nederlandse, Akademie van We- 
temschappen, Afd. Natuurkunde 53(1): 1—209. 

. 1970. Lepidoptera Tortricoidea from Tsaratan- 

ana Range [North Madagascar]. Mémoires 

O.R.S.T.O.M. 37: 103-150. 

. 1973. Tortricidae of the Andringitra Range, 

Central Madagascar (Lepidoptera). Part 1. Tortri- 

cinae. Bulletin de Muséum National d’ Historie 

Naturelle, no. 108, Zoologie 82: 105-143. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


. 1981. Tortricidae from Madagascar, Part 2. 
Olethreutinae, 1. Annales de la Société Entomo- 
logique de France (new series) 17: 7—32. 

. 1983. Tortricidae from Madagascar. Part 2. 
Olethreutinae, 2. Annales de la Société Entomo- 
logique de France (new series) 19: 291—310. 

Heppner, J. B. 1991. Faunal regions and the diversity 
of Lepidoptera. Tropical Lepidoptera (Gaines- 
ville), Vol. 2, supplement 1. 85 pp. 

. 1996. Keys to Families of Lepidoptera. Trop- 
ical Lepidoptera (Gainesville), Vol. 4, supplement 
3. 28 pp. 

Epstein, M. E. 1996. Revision and Phylogeny of the 
Limacodid-group Families, with Evolutionary 
Studies on Slug Caterpillars (Lepidoptera: Zyg- 
aenoidea). Smithsonian Contributions to Zoology 
582. 102 pp. 

Kristensen, N. P., ed. 1999. Handbook of Zoology, Vol. 
IV Arthropoda: Insects, Part 35 Lepidoptera, 
Moths and Butterflies. Walter de Gruyter, Berlin. 


John W. Brown, Systematic Entomology 
Laboratory, PSI, Agricultural Research 
Service, U.S. Department of Agriculture, 
c/o National Museum of Natural History, 
Washington, DC 20560-0168, U.S.A. (e- 
mail: jbrown@ sel.barc.usda.gov) 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 1077-1087 


OBITUARY 


Clyde Fuhriman Smith 
(1913-2000) 


Clyde Fuhriman Smith enjoyed a distin- 
guished, multi-faceted career in entomolo- 
gy, with an enduring interest in the taxon- 
omy of aphids and their host-plant associ- 
ations. He was born 10 August 1913, in 
Franklin County, Idaho, where his early ed- 
ucation began. Smith went on to study en- 
tomology, earning degrees at Utah State 
Agricultural College (now Utah State Uni- 
versity: B.S. 1935, M.S. 1938: George FE 
Knowlton, major advisor) and the Ohio 
State University (Ph.D. 1939: Alvah Peter- 
son, major advisor). His entomological ed- 


ucation extended far beyond course work 
with part-time or short, full-time periods as 
research assistant, teaching assistant, exten- 
sion assistant, and research fellow, expos- 
ing him to a wide range of insects and 
crops. 

In September 1939, Smith accepted a po- 
sition in Raleigh as an Assistant Entomol- 
ogist at the North Carolina Agricultural Ex- 
periment Station and N.C. State College 
(now University) and subsequently pro- 
gressed to Associate Professor (1943- 
1950), Professor and Department Head 


1078 


(1951-1963), Professor (1964-1979), and 
Professor Emeritus (1980—2000). Smith 
worked on insects and mites associated 
with nearly every major crop plant in North 
Carolina as documented by his publications 
(listed below). Under his leadership, the 
Department of Entomology (initially a sec- 
tion of the Division of Biological Sciences) 
more than doubled in number of faculty. 
After his tenure as head, he returned to re- 
search on cucumber and peach insects, no- 
tably the management of the pickleworm, a 
complex problem requiring intensive and 
complete control. As a professor emeritus, 
he continued publishing on aphid system- 
atics and served on the NCSU Insect Col- 
lection’s Museum Council. 

Stretching far beyond his microscope and 
boxes of prepared specimens, Smith’s lab- 
oratory embraced North Carolina’s or- 
chards, forests, farms, gardens, pest-infest- 
ed buildings, and roadside vegetation. He 
excelled at working with and gaining the 
trust of farmers and orchardists, talents vital 
to conducting meaningful scientific re- 
search in the days before university-sup- 
ported experimental research farms. 

Smith had an extraordinary ability to 
bring together scientific colleagues and oth- 
ers to exchange information. The present 
names of organizations he founded or 
helped to found are the Southeastern Peach 
Workers Conference (initiated 1948), N.C. 
Agricultural Chemicals School (1949), N.C. 
Pest Control Technicians School (1950), 
N.C. Pest Control Association (1950), N.C. 
Entomological Society (1956, first Presi- 
dent), and Heliothis Conference (late 1950s, 
now meeting with the Entomological So- 
ciety of America). In 1972, Smith received 
the N.C. Entomological Society’s Entomol- 
ogist of the Year award. Furthermore, he 
held membership in several other scholarly 
societies: Entomological Society of Amer- 
ica (President of Southeastern Branch in 
1972), Entomological Society of Washing- 
ton, Society of Systematic Zoology, Geor- 
gia Entomological Society, Gamma Sigma 
Delta, Society of the Sigma Xi, and Phi 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Kappa Phi. Smith was also instrumental in 
the passage of the N.C. Structural Pest Con- 
trol Act of 1955 and served as chair of the 
N.C. Structural Pest Control Commission 
from 1955 to 1967. 

Internationally known as a leading re- 
searcher on the systematics of aphids and 
their parasites, Smith deposited his large re- 
search collection of aphids at the NCSU In- 
sect Collection. It is rich in material from 
North Carolina and Utah, but includes spec- 
imens from around the world as well as 
Smith’s collection notebooks (including 
field notes). His collection of small parasitic 
wasps (Hymenoptera: Braconidae: Aphidi- 
inae) is also held at NC State. Additionally, 
the W. FE Barr Entomological Collection, 
University of Idaho, Moscow, and the En- 
tomological Museum of Utah State Univer- 
sity, Logan, both hold significant numbers 
of aphid slides donated by Smith. The pri- 
mary types for most (75) of the aphid spe- 
cies that Smith authored (or coauthored) are 
deposited at the U.S. National Collection of 
Insects and Mites housed at Beltsville, 
Maryland (www.sel.barc.usda.gov/aphid/ 
aphframe.htm). 

Among Smith’s numerous publications 
on aphids are: Aphididae of Puerto Rico 
(1963, with L. F Martorell and M. E. Pérez- 
Escolar), Bibliography of the Aphididae of 
the World (1972b), Keys to and Descrip- 
tions of the Genera of Pemphigini in North 
America (1974), An Annotated List of Aphi- 
didae of North America (1978, with C. S. 
Parron), An Annotated List of Aphididae of 
the Caribbean Islands and South America 
(1979, with M. M. Cermeli [Cermeli-Lol- 
lini]), and A Key to Many of the Common 
Alate Aphids of North Carolina (1992, with 
R. W. Eckel and E. Lampert). Smith’s doc- 
toral dissertation, which treated a group of 
small wasps parasitic on aphids, was the ba- 
sis for his Aphidiinae of North America 
(1944c). 

The extensive collection of papers Smith 
used to compile his bibliographies, check- 
lists and taxonomic works is deposited with 
the NCSU Libraries’s Special Collections 


VOLUME 102, NUMBER 4 


Department as the Clyde FE Smith Papers 
(Collection Number MC 221). These pa- 
pers, some of which are rare, encompass 
literature, reprints, pamphlets, and books, 
primarily pertaining to the insect family 
Aphididae. Associated with the papers are 
approximately 288,000 index cards on 
Aphididae with indexes that include: an an- 
notated generic and specific index to all lit- 
erature cited in Smith’s 1972 Bibliography 
of Aphididae, a host-plant index indicating 
the aphids associated with each host, the 
source citing this association, and an index 
of aphid-parasitoid associations. Together, 
the 1972 Bibliography and the 1978 An- 
notated List of Aphididae of North America 
provide indexes to the Smith Papers. 

His hobbies included gardening, hunting, 
fishing, lapidary, and genealogical research. 
Another more unusual pastime was watch- 
ing Hopkins’ Bioclimatic Law play itself 
out among the plants and associated arthro- 
pods of North Carolina, a state with ex- 
tremes in topography. This empirical law, 
which Smith taught in his fruit insect 
course, states that: 


“Other conditions being equal, the 
variation in the time of occurrence of 
a given periodical event in life activity 
in temperate North America is at the 
general average rate of 4 days to each 
I degree of latitude, 5 degrees of lon- 
gitude and 400 feet of altitude, later 
northward, eastward and upward in 
the spring and early summer, and the 
reverse in late summer and autumn.” 
(Hopkins, A. D. 1918. United States 
Department of Agriculture, Monthly 
Weather Review Supplement 9: 7). 


Over a 10-year period, Smith observed the 
seasonal development of a scrawny flow- 
ering peach tree near his office. He used it 
as an index to the first appearance in the 
spring of the adults of the plum curculio, 
whose grubs would later infest the peaches 
of the North Carolina Sandhills. Smith had, 
of course, observed and predicted other 
phenological events across the state corre- 


1079 


lated with the vernal development of that 
peach tree. He then would cajole his grad- 
uate students into bets of soft drinks con- 
cerning what they would find at the re- 
search site when they arrived. Smith lost a 
few sodas, but not many. 

Frequently, he suggested graduate stu- 
dent research projects that ranged far from 
his own expertise, expecting and inspiring 
students to learn on their own, as he had 
throughout his life. His graduate students 
included: R. Bastida, C. S. Black, Jr, C. S. 
Parron, and L. Van Balen (all M.S.); M. M. 
Cermeli-Lollini, H. L. Comroy, M. H. Far- 
rier, J. Graham, H. B. Moore, Jr., A. T. Ol- 
ive, M. E. Pérez-Escolar, and C. G. Wright 
(all Ph.D.). Smith genuinely enjoyed ob- 
serving and puzzling out nature, including 
the complex life histories of his beloved 
aphids. 

On 13 February 2000, his long, produc- 
tive life drew to a close, following a period 
of ill-health initiated by a stroke in January 
1999. 

Clyde FE Smith will be long remembered 
for his devotion to family, community, and 
church (Church of Jesus Christ of Latter 
Day Saints) as well as his professional 
achievements. He will be missed by his 
many friends, co-workers, and family mem- 
bers, particularly his ever supportive wife, 
Crystle; three children, Clara Beth, Caro- 
lynn, and Clyde Leslie, and their spouses; 
15 grandchildren; and 14 great-grandchil- 
dren. 


PUBLICATIONS BY CLYDE F. SMITH! 


Knowlton, G. F, J. S. Stanford, and C. F 
Smith. 1934. Birds as predators of the 
beet leafhopper. Journal of Economic 
Entomology 27: 1196—1197. 

Knowlton, G. FE and C. E Smith. 1935a. To- 


! Dr. Smith maintained a card index of his publica- 
tions with each item numbered. The 156 items in that 
index, plus nine others, are listed below, as far as pos- 
sible following Smith’s numerical ordering. Our efforts 
to verify the citations of certain older Extension pub- 
lications and a movie have so far proven unsuccessful. 
Two further papers are in review. 


1080 


mato fruitworms. Utah State Agricultural 
Experiment Station Leaflet 61: 1—4. 

Knowlton, G. FEF and C. E Smith. 1935b. 
The desert gridiron-tailed lizard. Copeia 
2: 102-103. 

Knowlton, G. E and C. E Smith. 1935c. 
Toads in the control of auto-camp in- 
sects. Journal of Economic Entomology 
28: 496. 

Knowlton, G. EF and C. FE Smith. 1935d. 
Currant and gooseberry aphids. Utah 
State Agricultural Experiment Station 
Leaflet 63: [1—4]. 

Knowlton, G. FE and C. E Smith. 1935e. 
Beet leafhopper predators—Birds. Pro- 
ceedings of the Utah Academy of Sci- 
ences, Arts, and Letters 12: 249-253. 

Knowlton, G. FE and C. E Smith. 1935f. 
Notes on Utah Scarabaeidae and Chry- 
somelidae (Coleoptera). Entomological 
News 46: 241-244. 

Knowlton, G. FE and C. E Smith. 1936a. 
Lizard aids in range insect control. Jour- 
nal of Economic Entomology 29: 461. 

Knowlton, G. EF and C. E Smith. 1936b. 
The aphid genus Epameibaphis in Utah. 
Proceedings of the Entomological So- 
ciety of Washington 38: 89-92. 

Knowlton, G. EF and C. E Smith. 1936c. 
Capitophorus aphids infesting Chryso- 
thamnus. Canadian Entomologist 68: 
107-113. 

Knowlton, G. E and C. E Smith. 1936d. 
Notes on intermountain aphids. Ento- 
mological News 47: 210-213. 

Knowlton, G. E and C. EF Smith. 1936e. 
Rose insects. Proceedings of the Utah 
Academy of Sciences, Arts, and Letters 
13: 263-267. 

Knowlton, G. E and C. E Smith. 1936f. 
Strawberry insects. Proceedings of the 
Utah Academy of Sciences, Arts, and 
Letters 13: 289-292. 

Knowlton, G. EK and C. E Smith. 1936g. 
Capitophorus aphids infesting Artemi- 
sia. Canadian Entomologist 68: 229— 
234. 

Knowlton, G. EK and C. E Smith. 1936h. 
Descriptions of some rabbit brush and 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


willow aphids. Annals of the Entomo- 
logical Society of America 29: 776— 
778. 

Knowlton, G. E and C. FE Smith. 19361. 
Aphid studies. Festschrift zum 60. Ge- 
burtstage von Professor Dr. Embrik 
Strand (Riga, Latvija) 1: 487—492, plate 
VIII. 

Smith, C. EF 1937. The aphid genus Fla- 
bellomicrosiphum in Utah. Pan-Pacific 
Entomologist 13: 127-129. 

Smith, C. E and G. E Knowlton. 1937. Ma- 
crosiphum aphids infesting Chryso- 
thamnus and Gutierrezia. Canadian En- 
tomologist 69: 269-272. 

Knowlton, G. EF and C. E Smith. 1937. 
Some aphids of the genus Capitopho- 
rus. Canadian Entomologist 69: 150— 
152: 

Smith, C. FE and G. EF Knowlton. 1938. The 
aphid genus Microsiphum in Utah and 
Idaho. Annals of the Entomological So- 
ciety of America 31: 162-166. 

Knowlton, G. FE and C. EF Smith. 1938a. 
Notes on western conifer aphids (Ho- 
moptera: Aphididae). Entomological 
News 49: 65-69. 

Knowlton, G. E and C. E Smith. 1938b. 
The aphid genus Pseudoepameibaphis. 
Journal of the New York Entomological 
Society 46: 217-222. 

Knowlton, G. F, C. E Smith, and FE C. 
Harmston. 1938. Pea aphid investiga- 
tions. Proceedings of the Utah Academy 
of Sciences, Arts, and Letters 15: 71— 
80. 

Smith, C. EF 1939a. The digestive system of 
Macrosiphum solanifolii (Ash.) (Aphi- 
dae: Homoptera). Ohio Journal of Sci- 
ence 39: 57-59. 

Smith, C. E 1939b. Hymenopterous para- 
sites of aphids with special reference to 
the Aphidiinae. Abstracts of Doctoral 
Dissertations (Ohio State University, 
Columbus) 30: 169-174. 

Smith, C. FEF and G. FE Knowlton. 1939. 
Three intermountain aphids. Canadian 
Entomologist 71: 241-243. 

Smith, C. E 1940. Notes on some Ohio 


VOLUME 102, NUMBER 4 


aphids. Ohio Journal of Science 40: 
139-142, plate I. 

Smith, C. E and G. E Knowlton. 1940. 
Three aphids of the genus Brevicoryne 
van der Goot. Annals of the Entomo- 
logical Society of America 33: 404— 
405. 

Smith, C. EF 194la. Control studies of the 
woolly apple aphid. Journal of Econom- 
ic Entomology 34: 590. 

Smith, C. E 1941b. A new species of hy- 
menopterous parasite of the pea aphid 
(Macrosiphum pisi Kaltentbach). An- 
nals of the Entomological Society of 
America 34: 537-538. 

Smith, C. FE 1941c. The genus Drepanaphis 
Del Guercio east of the Rocky Moun- 
tains. Journal of the Elisha Mitchell So- 
ciety 57: 226—242. 

Smith, C. E 1941d. Trichogramma and the 
Oriental fruit moth. Journal of Econom- 
ic Entomology 34: 590. 

Smith, C. E 1942. The use of cyanide in 
controlling the root form of the woolly 
apple aphid. Journal of Economic En- 
tomology 35: 908-910. 

Smith, C. E 1943a. North Carolina, p. 6. In 
State by State Summary of 1943 Or- 
chard Pest Problems. American Fruit 
Grower 63(2): 6. 

Smith, C. E 1943b. Scale insects harm 
peach trees in North Carolina. Research 
and Farming (North Carolina Agricul- 
tural Experiment Station, Raleigh) 1(2): 
12: 

Smith, C. EF 1943c. Experiments with the 
peach tree borer in North Carolina. 
Journal of Economic Entomology 36: 
215-218. 

Smith, C. E 1943d. Peach tree borer control 
recommendations, 1943. North Carolina 
Agricultural Experiment Station, 1943. 
[mimeograph printing] 

Smith, C. E and G. FE Knowlton. 1943. The 
aphid genus Drepanaphis Del Guercio. 
Journal of the Elisha Mitchell Society 
59: 171-176, plate 22. 

Smith, C. EF 1944a. Drepanaphis tissoti, a 


1081 


new species of aphid from Florida. Flor- 
ida Entomologist 27: [55]-57. 

Smith, C. E 1944b. Peach insect control. 
Proceedings of the Southern Workers’ 
Conference on Entomology (18th An- 
nual Meeting, February 1—3, 1944) 18: 
63-67. 

Smith, C. E 1944c. The Aphidiinae of 
North America (Braconidae: Hymenop- 
tera). Ohio State University Press, Co- 
lumbus, xii + 154 pp. 

Smith, C. E 1944d. Peach tree borer, control 
recommendations for North Carolina. 
[alternate title: The peach tree borer in 
North Carolina]. North Carolina Agri- 
cultural Extension Circular 277: [1]-8. 

Smith, C. E 1944e. Controlling the plum 
curculio on peaches. Research and 
Farming (North Carolina Agricultural 
Experiment Station, Raleigh) 2(2): 2-3. 

Smith, C. E 1945. Reduced concentrations 
of ethylene dichloride for peachtree bor- 
er control. Journal of Economic Ento- 
mology 38: 500-501. 

Smith, C. E 1946a. Curculio control rec- 
ommendations for North Carolina. 
North Carolina State College, January 
14, 1946. [lithograph printing] 

Smith, C. FE 1946b. Controlling soybean in- 
sects. Research and Farming (North 
Carolina Agricultural Experiment Sta- 
tion, Raleigh) 4(3): 7, 9. 

Smith, C. E 1946c. Plum curculio, post har- 
vest spraying. North Carolina Agricul- 
tural Experiment Station, August 1, 
1946. [mimeograph printing] 

Smith, C. E 1946d. Peach insects, North 
Carolina—1946. Proceedings of the 
Cumberland-Shenandoah Fruit Work- 
ers’ Conference (23rd Annual Meeting) 
239) 20" 0r22 1: 

Smith, C. E 1947a. Controlling pea aphids. 
North Carolina Agricultural Extension 
Service, March 1947. [mimeograph 
printing] 

Smith, C. E 1947b. Burning to control the 
curculio. North Carolina Agricultural 
Experiment Station, March 12, 1947. 
[mimeograph printing] 


1082 


Smith, C. EF 1947c. Emergence of the plum 
curculio. North Carolina Agricultural 
Experiment Station, March 20, 1947. 
[mimeograph printing] 

Smith, C. E 1947d. Curculio emergence 
and petal fall spray. North Carolina Ag- 
ricultural Experiment Station, April 7, 
1947. [mimeograph printing] 

Smith, C. EF 1947e. Emergence of the plum 
curculio. North Carolina Agricultural 
Experiment Station, April 16, 1947. 
[mimeograph printing] 

Smith, C. E 1947f. Wormy drops. North 
Carolina Agricultural Experiment Sta- 
tion, April 28, 1947. [mimeograph 
printing] 

Smith, C. EF 1947g. Notes on hexaethyl te- 
trophosphate. North Carolina Agricul- 
tural Experiment Station, June 3, 1947. 
[mimeograph printing] 

Smith, C. FE and C. N. Clayton. 1947. First 
generation curculio and brown rot con- 
trol. North Carolina Agricultural Exper- 
imental Station, June 9, 1947. [mimeo- 
graph printing] 

Smith, C. F 1947h. Plum curculio; post har- 
vest spraying. North Carolina Agricul- 
tural Experiment Station, August 19, 
1947. [mimeograph printing] 

Smith, C. FE 19471. Things to do and not to 
do; peach borer control; scale insects 
and burning. North Carolina Agricultur- 
al Experiment Station, September 1947. 
[mimeograph printing] 

Clayton, C. N. and C. FE Smith. 1947. Apple 
spray suggestions for 1947. North Car- 
olina Agricultural Experiment Station, 
1947. [mimeograph printing] 

Smith, C. FE and O. Veerhoff. 1947. Ethyl- 
ene dichloride injury to peach. Journal 
of Economic Entomology 40: 588. 

Smith, C. FE and W. H. Rankin. 1947. Does 
fertilization increase grain aphids? Re- 
search and Farming (North Carolina 
Agricultural Experiment Station, Ra- 
leigh) 6(1): 6. 

Smith, C. FE 1947}. Plum curculio control in 
North Carolina—1947. Proceedings of 
the | Cumberland-Shenandoah _ Fruit 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Workers’ Conference (24th Annual 
Meeting) 24: 26-29. 

Smith, C. EF and C. N. Clayton. 1948. Peach 
spray information for 1948. North Car- 
olina Agricultural Experiment Station 
Special Circular 5. [pagination un- 
known] 

Clayton, C. N. and C. F Smith. 1948. Apple 
spray information for 1948. North Car- 
olina Agricultural Experiment Station 
Special Circular 6. [pagination un- 
known] 

Smith, C. E 1948a. Pea aphid control in 
North Carolina. North Carolina Agri- 
cultural Experiment Station Special Cir- 
cular 7: [1-7]. 

Smith, C. EF 1948b. A new aphid on sweet 
potato. Florida Entomologist 31: 24-26. 

Smith, C. E 1948c. Plum curculio control 
in North Carolina. Journal of Economic 
Entomology 41: 220-227. 

Smith, C. EF 1948d. A new aphid on devil 
shoe string (Tephrosia virginiana, L. 
Pers.). Annals of the Entomological So- 
ciety of America 41: 384-386. 

Smith, C. F and C. N. Clayton. 1949. Peach 
spray information for 1949. North Car- 
olina Agricultural Experiment Station 
Special Circular 5(revised): 1-8. 

Clayton, C. N. and C. EF Smith. 1949. Apple 
spray information for 1949. North Car- 
olina Agricultural Experiment Station 
Special Circular 6(revised): 1-8. 

Smith, C. F, I. D. Jones, and J. A. Rigney. 
1949. Effect of insecticides on the fla- 
vor of peaches—1948. Journal of Eco- 
nomic Entomology 42: 618-623. 

Smith, C. E 1949. DDT-parathion: new 
weapon for peach growers. Research and 
Farming (North Carolina Agricultural 
Experiment Station, Raleigh) 7(3): 3. 

Smith, C. FE and C. Black. 1950. Peach bor- 
er paradox. Research and Farming 
(North Carolina Agricultural Experi- 
ment Station, Raleigh) 8(3): 5, 7. 

Smith, C. FE, I. D. Jones, and L. D. Calvin. 
1950. Effect of insecticides on the fla- 
vor of peaches—1949. Journal of Eco- 
nomic Entomology 43: 179-181. 


VOLUME 102, NUMBER 4 


Smith, C. FE and C. N. Clayton. 1950. Peach 
spray information. North Carolina Ag- 
ricultural Experiment Station Special 
Circular 5(revised): 1-[11]. 

Clayton, C. N. and C. EK Smith. 1950. Apple 
spray information. North Carolina Ag- 
ricultural Experiment Station Circular 
6(revised): 1—12. 

Randall, G. O., H. R. Garriss, and C. E 
Smith. 1951. Successful rose culture. 
North Carolina Agricultural Extension 
Service, Extension Circular 200(re- 
vised): [1—26]. 

Smith, C. E and C. N. Clayton. 1951. Peach 
spray information. North Carolina Ag- 
ricultural Experiment Station Special 
Circular 12: 1-11. 

Smith, C. E and L. Bennett. 1951. Some 
tobacco insects and their control. Mov- 
ie, 16 mm., color, sound. North Carolina 
State College, Raleigh, December 1951. 

Smith, C. E and C. N. Clayton. 1953a. 
Peach spray information, 1953. North 
Carolina Agricultural Experiment Sta- 
tion Special Circular 17: 1-11. 

smith, (E> By and'C, N. Clayton: 1953b. 
Peach spray information, 1953. North 
Carolina Agricultural College Plant Pa- 
thology Information Note 31: 10—15. 

Turnipseed,.G., FE. and ‘C.F Smith. 1953. 
Life history and control of scales on ap- 
ples in North Carolina. Journal of Eco- 
nomic Entomology 46: 969-972. 

Smith, C. EF, G. D. Jones, and R. L. Rabb. 
1953. Tobacco insect control. North 
Carolina Agricultural Extension Ser- 
vice, Raleigh, 4 pp. 

Smith, C. FE and C. N. Clayton. 1954. Peach 
spray information. North Carolina Ag- 
ricultural Experiment Station Special 
Circular 20: 1—12. 

Smith, C. EF and R. L. Rabb. 1954. The ef- 
fects of insecticides on the flavor of to- 
bacco. Proceedings, Association of 
Southern Agricultural Workers (51st 
Annual Convention, Dallas, Texas, Feb- 
ruary 1—3, 1954) 51: 107. 

Randall, G. O., H. R. Garriss, and C. FE 
Smith. 1954. Successful rose culture. 


1083 


North Carolina Agricultural Extension 
Circular 200(revised): [1—24]. 

Clayton, C. N., H. C. Fink, C. EK Smith, and 
G. E Turnipseed. 1954. Apple spray in- 
formation. North Carolina Agricultural 
Experiment Station Special Circular 19: 
1-16. 

Rabb, R. L., E E. Guthrie, H. E. Scott, and 
C. E Smith. 1955. Tobacco insects of 
North Carolina and their natural ene- 
mies. North Carolina Agricultural Ex- 
periment Station Bulletin 394: [1]-35. 

Harris, J. H., E A. Hassis [Haasis], and C. 
FE Smith. 1955. Azaleas and camellias. 
North Carolina Agricultural Extension 
Circular 246: [1-32]. 

Clayton, C. N. and C. FE Smith. 1956. Apple 
disease and insect control recommen- 
dations—1956. North Carolina Pesti- 
cide Manual 1956: 27-28. 

Smith, C. FE and C. N. Clayton. 1956. Peach 
insect and disease control recommen- 
dations—1956. North Carolina Pesti- 
cide Manual 1956: 29-31. 

Smith, C. EF 1956. Zeno Payne Metcalf, dis- 
tinguished entomologist. Science 123: 
1022. 

Metcalf, Z. P. and C. EK Smith. 1956. Intro- 
duction, pp. ii—vil. Jn Metcalf, Z. P, 
ed., General Catalogue of the Homop- 
tera. Fascicle IV. Fulgoroidea. Part 18. 
Eurybrachidae and Gengidae. North 
Carolina State College, Raleigh. 

Metcalf, Z. P and C. E Smith. 1957. Intro- 
duction, pp. ui—vill. In Metcalf, Z. P,, 
ed., General Catalogue of the Homop- 
tera. Fascicle IV. Fulgoroidea. Part 13. 
Flatidae and Hypochthonellidae. North 
Carolina State College, Raleigh. 

Clayton, C. N., G. E Turnipseed, and C. E 
Smith. 1957. Apple spray information. 
North Carolina Extension Circular 406: 
1-20. 

Smith, C. EK and C. N. Clayton. 1957. Peach 
spray information. North Carolina Ex- 
tension Circular 407: 1—16. 

Harris, J. H., E A. Hassis [Haasis], and C. 
E Smith. 1957. Azaleas and camellias. 


1084 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


North Carolina Agricultural Extension 
Circular 246 (revised): [1—32]. 

Metcalf, Z. P. and C. EF Smith. 1958. Intro- 
duction, pp. ili-vu. In Metcalf, Z. P, 
ed., General Catalogue of the Homop- 
tera. Fascicle IV. Fulgoroidea. Part 15. 
Issidae. North Carolina State College, 
Raleigh. 

Smith; <G) B.-L: EF Martorell;and*M~. E: 
Pérez-Escolar. 1958. Myzus persicae 
(Sulzer) in Puerto Rico. Journal of Ag- 
riculture of the University of Puerto 
Rico 42: 263-266. 

Smith, C. EF 1958. What’s new in insecti- 
cides? Agricultural Chemicals 13(3): 
43, 45. 

Smith, C. EF 1959. A new species of aphid 
on sugar maple (Aphidae: Homoptera). 
Annals of the Entomological Society of 
America 52: 647-649. 

Smith, C. FE and C. N. Clayton. 1959. Peach 
spray information. North Carolina Ag- 
ricultural Extension Circular 407(tre- 
vised): 1-15. 

Smith, C. EF 1960a. Aphids (Aphidae: Ho- 
moptera) on “‘cacao”’ in the Dominican 
Republic. Journal of Agriculture of the 
University of Puerto Rico 44: 154-156. 

Smith, C. E 1960b. New species of Aphi- 
dae: Homoptera from Puerto Rico. Jour- 
nal of Agriculture of the University of 
Puerto Rico 44: 157-162. 

Smith, C. FE and N. Tuatay. 1960. The genus 
Microparsus Patch (Aphidae: Homop- 
tera). Annals of the Entomological So- 
ciety of America 53: 735-742. 

Smith, C. EF 1960c. North Carolina finds 3- 
year old inspection form still accept- 
able. Pest Control 28(9): 40—41. 

Harris, J. H., EF A. Haasis, and C. E Smith. 
1960a. Azaleas and camellias. North 
Carolina Agricultural Extension Circu- 
lar 246(revised): [1—32]. 

Harris, J. H., E A. Haasis, and C. E Smith. 
1960b. How to grow azaleas and ca- 
mellias. North Carolina Agricultural 
Extension Service, Extension folder. 

Smith, C. E 1961. p.194. In Comments of 
the proposed use of the plenary powers 


to designate a type-species for Eucera- 
phis Walker, 1870. Z.N.(S.) 1363. Bul- 
letin of Zoological Nomenclature 18: 
194. 

Smith, C. FE 1962. p. 196. In Discussion of 
the case “Aphis Linnaeus, 1758, its 
type-species, and the family-group 
name derived from it (Insecta: Hemip- 
tera). Z.N.(S) 881”. Bulletin of Zoolog- 
ical Nomenclature 19: 195-198. 

Smith, C. E and O. E. Heie. 1963. Me- 
gouroparsus, new genus, related to Mi- 
croparsus Patch and Megoura Buckton 
(Homoptera: Aphidae). Annals of the 
Entomological Society of America 56: 
401—406. 

Fields, H. M., G. C. Klingman, C. FE Smith, 
and C. N. Clayton. 1963. Pesticides, 
facts on their use in North Carolina. Na- 
tional Agricultural Chemicals Associa- 
tion News and Pesticide Review 21(4): 
8—10. 

Smith, C. E, L. E Martorell, and M. E. 
Pérez-Escolar. 1963. Aphididae of 
Puerto Rico. University of Puerto Rico 
Agricultural Experiment Station Tech- 
nical Paper 37: [1]-121. 

Smith, C. F 1965a. Tiliphagus lycoposugus, 
new genus, new species (Aphididae: 
Homoptera) from Tilia americana and 
Lycopus virginicus. Annals of the En- 
tomological Society of America 58: 
781-786. 

Smith, C. F 1965b. Book review: Ento- 
mological techniques. How to work 
with insects, by Alvah Peterson. Bulle- 
tin of the Entomological Society of 
America 11: 96. 

Smith, C. E and G. EK Knowlton. 1966. The 
genus Aspidaphium Borner (Homop- 
tera: Aphididae). Pan-Pacific Entomol- 
ogist 42: 20-24. 

Smith, C. FE and J. Graham. 1967. Life his- 
tory, synonymy, and description of Neo- 
prociphilus aceris (Homoptera: Aphi- 
didae). Annals of the Entomological So- 
ciety of America 60: 67-72. 

Smith, C. F and D. G. Dillery. 1968. The 
genus Drepanaphis Del Guercio (Ho- 


VOLUME 102, NUMBER 4 


moptera: Aphididae). Annals of the En- 
tomological Society of America 61: 
185-204. 

Smith, C. E and J. O. Pepper. 1968. Gryl- 
loprociphilus frosti, new genus, new 
species, from the eastern United States 
(Homoptera: Aphididae). Proceedings 
of the Entomological Society of Wash- 
ington 70: 57—60. 

Clayton, C. N., G. T. Weekman, and C. E 
Smith. 1968. Peach spray program. 
North Carolina Pesticide Manual 1968: 
a6: 

Smith, C. EK 1969a. Pemphiginae associated 
with the roots of conifers in North 
America (Homoptera: Aphididae). An- 
nals of the Entomological Society of 
America 62: 1128-1152. 

Smith, C. E 1969b. Controlling peach scale. 
Research and Farming (North Carolina 
Agricultural Experiment Station, Ra- 
leigh) 28(1—2): 12. 

Smith, C. E and C. N. Clayton. 1970. Peach 
disease and insect control in North Car- 
olina. North Carolina Agricultural Ex- 
tension Circular 407: 1-17. 

Smith, C. FE 1970a. Emergence of the peach 
tree borer in North Carolina. Journal of 
Economic Entomology 63: 1700-1701. 

Smith, C. EF 1970b. Notes on the genus Pic- 
turaphis and related genera with a new 
species of Picturaphis from Puerto Rico 
(Aphididae: Homoptera). Journal of Ag- 
riculture of the University of Puerto 
Rico 54: 683-688. 

Clayton, C. N., C. E Smith, and G. T. 
Weekman. 1970. Peach spray program. 
North Carolina Agricultural Chemicals 
Manual 1970: 112-113. 

Smith, C. F 1970c. Sexual activity and egg 
fertility of the peach tree borer. Journal 
of Economic Entomology 63: 1909— 
1910. 

Clayton, C. N., C. E Smith, and K. A. Sor- 
ensen. 1971. Peach spray program. 
North Carolina Agricultural Chemicals 
Manual 1971: 124-125. 

Smith, C. F, S. M. Gaud, L. E Martorell, 
and M. E. Pérez-Escolar. 1971. Addi- 


1085 


tions and corrections to the Aphididae 
of Puerto Rico. Journal of Agriculture 
of the University of Puerto Rico 55: 
192-258. 

Smith, C. E and G. EF Knowlton. 1971. 
Pleotrichophorus tetradymiae, a new 
species of aphid from Utah (Homoptera: 
Aphididae). Proceedings of the Ento- 
mological Society of Washington 73: 
320-323. 

Smith, C. E 1971. The life cycle and re- 
description of Mordvilkoja vagabunda 
(Homoptera: Aphididae). Proceedings 
of the Entomological Society of Wash- 
ington 73: 359-367. 

Smith, C. FE and H. L. G. Stroyan. 1972. 
The probable identity of Prociphilus 
(Pulvius) probosceus (Homoptera: 
Aphididae). Annals of the Entomologi- 
cal Society of America 65: 804-807. 

Smith, C. E and C. N. Clayton. 1972. Peach 
disease and insect control in North Car- 
olina. North Carolina Agricultural Ex- 
tension Circular 407(revised): [1]-17. 

Clayton, C. N., C. EF Smith, and K. A. Sor- 
ensen. 1972. Peach spray program. 
North Carolina Agricultural Chemicals 
Manual 1972: 124-125. 

Smith, C. EF 1972a. Synonymy, redescrip- 
tion, and biology of Neoparacletus cor- 
rugatans (Homoptera: Aphididae). An- 
nals of the Entomological Society of 
America 65: 1328-1331. 

Smith, C. E 1972b. Bibliography of the 
Aphididae of the world. North Carolina 
Agricultural Experiment Station Tech- 
nical Bulletin 216: [1]-717. 

Clayton, C. N., C. E Smith, and K. A. Sor- 
ensen. 1973. Peach spray program. 
North Carolina Agricultural Chemicals 
Manual 1973: 221-222. 

Clayton, C. N., C. E Smith, and K. A. Sor- 
ensen. 1974. Peach spray program. 
North Carolina Agricultural Chemicals 
Manual 1974: 251-252. 

Smith, C. EF 1974a. The genus Phloeomyzus 
with the description of P. dearborni n. 
sp. from Populus tremuloides Michx. 
(Homoptera: Aphididae). Proceedings 


1086 


of the Entomological Society of Wash- 
ington 76: 66—72. 

Smith, C. E and S. M. Gaud. 1974. The 
alate viviparae of Picturaphis Blanchard 
(Homoptera: Aphididae). Journal of Ag- 
riculture of the University of Puerto 
Rico 58: 381-383. 

Smith, C. E 1974b. Keys to and descrip- 
tions of the genera of Pemphigini in 
North America (Homoptera: Aphididae: 
Pemphiginae). North Carolina Agricul- 
tural Experiment Station Technical Bul- 
letin 266: [i]—vi, [1]-61. 

Smith, C. EF and G. E Knowlton. 1975. 
Moss aphids in the United States (Ho- 
moptera: Aphididae). United States De- 
partment of Agriculture Cooperative 
Economic Insect Report 25: 423—431. 

Smith, C. FE and A. G. Robinson. 1975. The 
genus Myzodium with the description of 
M. knowltoni, new species (Homoptera: 
Aphididae). Proceedings of the Ento- 
mological Society of Washington 77: 
48 1—486. 

Clayton, C. N., C. E Smith, and K. A. Sor- 
ensen. 1975. Peach spray program. 
North Carolina Agricultural Chemicals 
Manual 1975: 251-252. 

Smith, C. E and C. N. Clayton. 1976a. 
Peach disease and insect control in 
North Carolina. North Carolina Agri- 
cultural Extension Circular 407(re- 
vised): 1-19. 

Stary, P. and C. E Smith. 1976. The Nearc- 
tic distribution, host plants and nomen- 
clature of Monoctonus (Homoptera: 
Aphididae). Proceedings of the Ento- 
mological Society of Washington 78: 
171-175. 

Smith, C. EF and C. N. Clayton. 1976b. 
Peach spray program. North Carolina 
Agricultural Extension Circular 407(sup- 
plement): 1—4. 

Smith, C. E and G. FE Knowlton. 1977. The 
genus Rhopalosiphoninus Baker (Ho- 
moptera: Aphididae) in North America. 
United States Department of Agricul- 
ture Cooperative Plant Pest Report 2: 
75-80. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Smith, C. E and C. N. Clayton. 1978a. 
Peach spray information. North Caroli- 
na Agricultural Extension Service Cir- 
cular 407(supplement, revised): [1—4 
(folded leaflet)]. 

Smith, C. E and C. N. Clayton. 1978b. 
Peach disease and insect control in 
North Carolina. North Carolina Agri- 
cultural Extension Service AG-146 [re- 
vision of former Circular 407]: [i—1i, 1]- 
YS) 

Smith, C. E and C. S. Parron. 1978. An 
annotated list of Aphididae (Homop- 
tera) of North America. North Carolina 
Agricultural Experiment Station Tech- 
nical Bulletin 255: [iJ—viii, 1—428. 

Smith, C. EF and W. V. Campbell. 1979. 
Charles Henry Brett 1909-1978. Jour- 
nal of Economic Entomology 72: 157. 

Smith, C. E and M. M. Cermeli [Cermeli- 
Lollini]. 1979. An annotated list of 
Aphididae (Homoptera) of the Carib- 
bean Islands and South and Central 
America. North Carolina Agricultural 
Research Service Technical Bulletin 
259: [iii], 1-131. 

Cermeli L[ollini], M. [M.], and C. E Smith. 
1979. Keys to species of the genus Pic- 
turaphis (Homoptera: Aphididae) with 
descriptions of new species. Proceed- 
ings of the Entomological Society of 
Washington 81: 611—620. 

Smith, C. E 1980a. A fourth species of Tox- 
opterella Hille Ris Lambers (Homop- 
tera: Aphididae) from North America 
with a key to species. Proceedings of 
the Entomological Society of Washing- 
ton 82: 276—283. 

Smith, C. E 1980b. Notes and keys to the 
species of Carolinaia (Homoptera: 
Aphididae). Proceedings of the Ento- 
mological Society of Washington 82: 
312-318. 

Smith, C. E 1982. A key to the species of 
Hyalomyzus (Homoptera: Aphididae) in 
North America, with description of a 
new species. Proceedings of the Ento- 
mological Society of Washington 84: 
325-331. 


VOLUME 102, NUMBER 4 


Smith, C. EK and H. A. Denmark. 1982. Tri- 
chosiphonaphis polygoni (van der Goot) 
(Homoptera: Aphididae), a genus and 
species new to the United States. Flor- 
ida Entomologist 65: 381-382. 

Smith, C. E 1983. A new species of aphid 
(Homoptera: Aphididae) from Potentil- 
la canadensis. Proceedings of the En- 
tomological Society of Washington 85: 
64-68. 

Smith, C. E and G. FE Knowlton. 1983. A 
key to the species of aphids (Homop- 
tera: Aphididae) on wild Geranium spp. 
in the United States, with description of 
a new species. Proceedings of the En- 
tomological Society of Washington 85: 
686-690. 

Smith, C. F and H. A. Denmark. 1984. Life 
history and synonymy of Grylloproci- 
philus imbricator (Fitch) (Homoptera: 
Aphididae). Florida Entomologist 67: 
430-434. 

Smith, C. E 1985. Pemphiginae in North 
America, pp. 277—302. In Szelegiewicz, 
H., ed., Evolution and Biosystematics of 
Aphids. Proceedings of the International 
Aphidological Symposium at Jablonna, 
5-11 April 1981. Zaklad Narodowy im 
Ossolinskich Wyawnictwo, Warsaw. 

Smith, C. FE, R. W. Eckel, and E. Lampert. 
1992. A key to many of the common 
alate aphids of North Carolina (Aphi- 
didae: Homoptera). North Carolina Ag- 
ricultural Research Service Technical 
Bulletin 299: [i]—vi, 1—92. 

Smith, C. E and R. W. Eckel. 1996. Aphis 
crassicauda n. sp. (Homoptera: Aphi- 


1087 


didae), with a key to the alate species 
of Aphis on Viburnum spp. in the Unit- 
ed States. Proceedings of the Entomo- 
logical Society of Washington 98: 44— 
49. 

Clements, K. M., B. M. Wiegmann, C. E. 
Sorenson, C. FE Smith, P. A. Neese, and 
R. M. Roe. 2000. Genetic variation in 
the Myzus persicae complex (Homop- 
tera: Aphididae): evidence for a single 
species. Annals of the Entomological 
Society of America 93: 31—46. 


ACKNOWLEDGMENTS 


For useful suggestions and information, 
we thank: Herbert H. Neunzig, Charles G. 
Wright (both Professors Emeriti), Carol S. 
Parron (former Curator of the NCSU Insect 
Collection), James D. Harper, John R. Mey- 
er, Michael R. Roe, and Clyde E. Sorenson 
(Department of Entomology, North Caroli- 
na State University, Raleigh); Caroline 
Weaver and Linda A. McCormick (Special 
Collections Department, North Carolina 
State University Libraries); Pamela E. Pur- 
year (Agricultural Communications—Tobac- 
co Literature Service, North Carolina State 
University); Randi V. W. Eckel (RVWE 
Consulting, Frenchtown, New Jersey); and 
Manya B. Stoetzel, (Systematic Entomolo- 
gy Laboratory, Agricultural Research Ser- 
vice, U.S. Department of Agriculture, 
Beltsville, Maryland). 


Maurice H. Farrier and Lewis L. Deitz, 
Department of Entomology, North Carolina 
State University, Raleigh, NC 27695-7613, 
U.S.A. (email: lewis.deitz@ ncsu.edu) 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 1088-1092 


SOCIETY MEETINGS 


1044th Regular Meeting—January 6, 2000 


The 1044th regular meeting of the En- 
tomological Society of Washington (ESW) 
was called to order in the Cathy Kerby con- 
ference room (CE-340) of the National Mu- 
seum of Natural History, Washington, D.C., 
by incoming President Dave Furth at 7:35 
pm. The meeting was attended by 20 mem- 
bers and 10 guests. Stu McKamey read the 
minutes of the 1043rd meeting, which were 
approved without modification. 

Chris Thompson called a point of order, 
that the ESW audit was not presented. Don 
Anderson reported that the audit was signed 
by committee members and provided a 
summary of the status: ESW is doing fine. 
Thompson moved to accept the report and 
the motion was quickly seconded and the 
audit approved unanimously. There were no 
new applicants for membership and no new 
members were present. One visitor was in- 
troduced. In miscellaneous business, there 
was short discussion about the ESW web- 
site, which has not yet materialized. 

For exhibits, Dave Furth announced the 
Washington Academy of Science website 
(http://www.washacadsci.org), displayed 
the calendars of the Entomological Society 
of America and the Organization for Trop- 
ical Studies, displayed insect toys recently 
collected without permits in Canada and 
Mexico, insect tacks (is nothing safe from 
the IRS?) and even a bee-motif shower cur- 
tain. Warren Steiner added to his ‘global 
warming’ insect sighting list: buckeye and 
sulpher butterflies spotted on December 12, 
to which a January 2nd monarch eclosion 
was added from the audience. Warren noted 
the insects included in the revamped 
NMNH elephant exhibit and lastly an- 
nounced a new publication: the Proceedings 
of the Appalachian Biogeography Sympo- 
sium, edited by ESW member Ralph Eck- 
erlin. Edd Barrows completed the exhibits 
with a butterfly mobile calendar. 


John Brown introduced the evening’s 
speaker, past ESW President Mike Schauff, 
who took us on a trip “Out and About 
Down Under: Collecting Tiny Wasps in 
Australia.’ In truth, three of his trips were 
cleverly merged into one tour, all beauti- 
fully documented with slides of cities, wa- 
terfalls, panoramic vistas, beaches, charis- 
matic megafauna, and not-so-charismatic 
megafauna: crocodiles, snakes, and huge 
leeches. We started our trip in subtropical 
SE Australia, which proved the best for his 
parasitic chalcidoid wasps. We traveled 
northward into the understory of the 
Queensland tropical rainforest, which was, 
as expected, poor collecting. From there we 
moved inland to the edge of the more arid 
central plateau, heavily agricultural but 
with open forests and drainage areas that 
yielded many chalcidoids. Then, drawn like 
a moth to flame, we headed into the ‘red 
center’ desert of the continent. Collecting 
was excellent in the shaded areas around 
Ayer’s Rock, Uluru National Park, where 
water persists year-round. We continued 
south to Adelaide, Bookmark Biosphere 
Reserve in the Murray River watershed, 
and Kangaroo Island, collecting wasps and 
enjoying the abundant marsupials, some- 
times scaring them away from the best 
shade spots for a break from the sun, other 
times sharing a research station with them. 
Eventually we had to return to Canberra, 
but couldn’t leave without collecting on 
Black Mountain, located just behind CSI- 
RO headquarters. Collecting permits for the 
trip were arranged by state and by National 
Park and required a couple months’ ad- 
vance planning. 

The meeting was adjourned at 8:50 pm. 
Refreshments were provided by John 
Brown. 


Respectfully submitted, 
Stuart H. McKamey 
Recording Secretary 


VOLUME 102, NUMBER 4 


1045th Regular Meeting—February 3, 
2000 


The 1045th regular meeting of the En- 
tomological Society of Washington (ESW) 
was called to order in the Cathy Kerby con- 
ference room (CE-340) of the National Mu- 
seum of Natural History, Washington, D.C., 
by President Dave Furth at 7:35 pm. The 
meeting was attended by 22 members and 
4 guests. Stu McKamey read the minutes of 
the 1044th meeting, which were approved 
with minor modification. 

Steve Lingafelter read the names of four 
new applicants for membership: Kevin 
O’Neill, Wayne Wehling, Qiao Gexia, and 
Daniel E. Perez-Gelabert. Two new mem- 
bers and two visitors were introduced. 

For exhibits, Chris Thompson called at- 
tention to a recent Nature article on the pu- 
tative forces driving the evolutionary origin 
of butterflies, an excellent new handbook 
for taxonomy (Describing Species: Practi- 
cal Taxonomic Procedure for Biologists), 
and a new, world catalog of beeflies. Edd 
Barrows noted a special issue of the Amer- 
ican Entomologist on biodiversity and had 
one on display. Ralph Eckerlin displayed a 
new Spanish language entomology text: Jn- 
sectos y Otros Artrépodos de Importancia 
Médica y Veterinaria, by E. Méndez. Dan- 
iel Perez had on hand two new publications 
from the Dominican Republic: a catalog of 
Dominican amber fossils and an Occasional 
Publication on taxonomy from the national 
museum. Dave displayed a recent Science 
article on ‘hire classification’ (i.e., charging 
for patronyms), a new curation book (Care 
and Conservation of Natural History Col- 
lections), a new book on the systematics of 
Western North American butterflies, and 
“Bug Bites,”’ chocolate candies with insect 
trading cards. 

Stu McKamey introduced the meeting’s 
speaker, Dr. Steven Lingafelter, ESW mem- 
ber and Research Scientist with the USDA 
Systematic Entomology Laboratory. He 
spoke to us about “‘Asian Longhorned Bee- 
tles of the Genus Anoplophora: a New 


1089 


Threat to the United States.”’ First detected 
in New York in 1996 but probably present 
for 10 years, these large, showy beetles 
have received wide media coverage owing 
to their destruction of living ornamental 
trees. The larvae are the real culprits, bor- 
ing in the live wood for up to a year. Con- 
trol here and even in Asia often consists of 
eradication: destroying any infested trees. 
Lingafelter’s connection to Anoplophora is 
his taxonomic expertise on longhorns (Cer- 
ambycidae) and the need to resolve species 
delimitations in the genus, with about 50 
nominal species, to aid control and inter- 
ception efforts. Additionally, all distribution 
records will be databased and an identifi- 
cation guide will be produced. Most records 
of the genus are from Japan, Korea, and 
China, where Lingafelter visited recently to 
examine institutional and private collec- 
tions, and to do more collecting. The Chi- 
nese portion was coordinated through the 
joint U.S.A.-China Institute of Biological 
Control and the Chinese Forestry Institute. 
The showy patches of color on the beetles’ 
elytra, traditionally used to delimit species, 
are actually quite variable among and even 
within populations. The genitalia are, sur- 
prisingly, not very useful either. The species 
appear to fall into five morphological 
groups defined by minor structural, external 
features. Quarantine controls have been 
strengthened and continued vigilance is re- 
quired to avoid additional introductions of 
Anoplophora. 

The meeting was adjourned at 8:53 pm. 
Refreshments were provided by John 
Brown. 


Respectfully submitted, 
Stuart H. McKamey 
Recording Secretary 


1046th Regular Meeting—March 2, 2000 


The 1046th regular meeting of the En- 
tomological Society of Washington (ESW) 
was called to order in the Cathy Kerby con- 
ference room (CE-340) of the National Mu- 
seum of Natural History, Washington, D.C., 


1090 


by President Dave Furth at 7:37 pm. The 
meeting was attended by 23 members and 
10 visitors. M. Alma Solis read the minutes 
of the 1045th meeting, which were ap- 
proved without modification. 

Steve Lingafelter read the names of three 
new applicants for membership: Ken Karns, 
Tam Nguyen, and Art Evans. One new 
member was introduced. 

For exhibits, Ralph Eckerlin brought the 
book entitled Kinabalu, edited by K. M. 
Wong & A. Phillipps and Chris Thompson 
brought Biology Systematics: Principles 
and Applications by Randall T. Schuh. 
Dave Furth brought the following books 
and articles: Ants at Work by Deborah Gor- 
don; Ecology of Insects: Concepts and Ap- 
plications, edited by M. R. Speight, M. D. 
Hunter & A. D. Watt; Forester Moths by 
K. A. Efetov & G. M. Tarmann; Insect 
Plant Relationships edited by S. J. Simp- 
son, A. J. Mordue & J. Hardie; The Johns 
Hopkins University of School of Hygiene 
and Public Health, Lloyd E. Rozeboom 
Mosquito Collection (1999. Amer. Mosqui- 
to Control Assoc. 15: 526-551), and two 
articles by Ashley Gurney entitled A Short 
History of the Entomological Society of 
Washington (1976. Proc. Entomol Soc. 
Wash. 78: 225-239), and Who Designed the 
Schmitt Box? (1975. Bull. Entomol. Soc. 
Amer. 21: 225-228). Edd Barrows brought 
some sawflies from Dyke Marsh. Gaby 
Chavarria advertised the film, Pollinators in 
Peril, to air on TBS Superstation, Monday, 
March 27, 11:00 pm/et., 8:00 pm/pt., and 
modeled a “‘pollinator’’ blouse worn by her. 

John Brown introduced the speaker, Dr. 
Gabriella Chavarria, National Fish and 
Wildlife Foundation (NWF) and Research 
Associate at the National Museum of Nat- 
ural History. Her talk was entitled ‘‘Polli- 
nation Conservation: the Truth About the 
Birds and the Bees.’ She spoke generally 
on pollination biology, flower morphology, 
kinds of bees, and co-evolution between 
flowers and pollinating organisms. At the 
NWFE she has been instrumental in a “Save 
the Pollinators”’ program. Every 3rd bite of 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


food is brought to humans by a pollinator. 
The best pollinators are bees (72.3%), 
among many others, including vertebrates 
such as birds and bats. But these pollinators 
are threatened by habitat destruction and by 
habitat and chemical habitat fragmentation. 
The major threats to Apis mellifera have 
been parasitic mites, bee diseases and pests, 
competition with africanized bees, and pes- 
ticide use. Fortunately, there are native bees 
to serve as alternative pollinators such as 
mason bees and bumblebees. Some solu- 
tions to the threats are: pollination research 
and monitoring, international agreements to 
protect migrating species, incorporation of 
wildlife corridors between agricultural are- 
as, environmentally safe golf courses, plant- 
ing native plants in backyards, and placing 
‘bee condominiums.”’ She suggested read- 
ing The Forgotten Pollinators by Steve 
Buckman & Gary Nabhan. 

The meeting was adjourned at 8:35 pm. 
Refreshments were provided by John 
Brown. 


Respectfully submitted, 
M. Alma Solis 


1047th Regular Meeting—April 6, 2000 


The 1047th regular meeting of the En- 
tomological Society of Washington (ESW) 
was called to order in the Cathy Kerby con- 
ference room (CE-340) of the National Mu- 
seum of Natural History, Washington, D.C., 
by President Dave Furth at 7:32 pm. The 
meeting was attended by 20 members and 
1 guest. Stu McKamey read the minutes of 
the 1046th meeting, which were approved 
with minor modification. 

Steve Lingafelter read the names of five 
new applicants for membership: Netta Dor- 
chin, James Kruse, Gordon E Pratt, Gale E. 
Ridge-O’Connor, and David Campbell. No 
new members were present. 

For exhibits, Ed Cohen displayed the No- 
vember 1999 issue of The American Mu- 
seum of Natural History containing an ar- 
ticle on the flight mechanics of insects. 
Dave Furth announced BugFest on the Mall 


VOLUME 102, NUMBER 4 


May 20th, encouraging ESW support; dis- 
played a new CD on the NHB East Court 
and the entomology compactor construc- 
tion; had the three latest issues of Food In- 
sects Newsletter from Montana State Uni- 
versity; announced CDs on beetles by John 
Lawrence, CSIRO; a world catalog of Cas- 
sidinae; The Cicadidae of China, in Chinese 
but full of photos [Ilustrataj Insect-faunoj: 
2, Chou & Lei (eds.)]; and a new Peterson 
field guide to western butterflies by P. Opler 
& A. Bartlett Wright. He also solicited 
opinions for an ESW website, should the 
project get started. Lastly, Furth announced 
the annual ESW banquet, tentatively sched- 
uled for June 8 with guest speaker Dave 
Grimaldi. 

John Brown introduced the night’s speak- 
er, Jim Whitfield (University of Arkansas/ 
University of Illinois) who presented his re- 
search on the ““Phylogeny of Microgastroid 
Braconid Wasps, and What it Tells Us 
About Polydnavirus Evolution.”’ Like other 
insects with endoparasitoid larvae, these 
wasp larvae battle the hosts’ immune sys- 
tems. Their special tools are polydnavirus- 
es: viral genes incorporated into their own 
genome and therefore inherited between 
generations. The expression of these genes 
suppresses the host’s immune response and 
can also alter its nutrition and developmen- 
tal rate. Dr. Whitfield investigated the origin 
of this virus-wasp relationship by compar- 
ing their phylogenies at three taxonomic 
levels: subfamilies, genera, and species. 
Looking at wasp subfamilies, evidence 
from morphology or molecules or both, and 
analyzed by parsimony or maximum like- 
lihood all confirmed a single origin of the 
association, with no subsequent loss. At the 
generic level, all data sets were congruent 
concerning which branches were short vs. 
long, but despite all attempts to wring out 
well supported hypotheses of relationship, 
all they could show was a rapid radiation 
of genera. That is, the problem seemed to 
be the evolutionary pace of the microgas- 
troid wasp genera, which is confirmed in- 
dependently by fossils, rather than flaws in 


1091 


the methods or data. At the species level, 
analysis is difficult because the viral genes 
are meshed into the wasp genome, yet sep- 
arated from each other. Whitfield and col- 
leagues so far have found two viral genes 
and their phylogenies match closely that of 
their wasp hosts. Dr. Whitfield concluded 
that the viral genes were probably incor- 
porated 60 to 80 million years ago and were 
followed by rapid diversification at the ge- 
neric level. 

The meeting was adjourned at 8:55 pm. 
Refreshments were provided by John 
Brown. 


Respectfully submitted, 
Stuart H. McKamey 
Recording Secretary 


1048th Regular Meeting—May 4, 2000 


The 1048th regular meeting of the En- 
tomological Society of Washington (ESW) 
was called to order in the Cathy Kerby con- 
ference room (CE-340) of the National Mu- 
seum of Natural History, Washington, D.C., 
by President Dave Furth at 7:36 pm. The 
meeting was attended by 18 members and 
4 guests. Stu McKamey read the minutes of 
the 1047th meeting, which were approved 
without modification. 

Steve Lingafelter, the membership chair, 
was in Nepal so no new applicants for 
membership were read. No new members 
were present. 

For exhibits, Warren Steiner passed 
around a live caterpillar hunter (Callasoma, 
Carabidae), John Brown announced the 
ESW Annual Banquet, June 8 at the Uni- 
formed Services University of Health Sci- 
ences in Bethesda, Maryland. The feature 
will be a talk about the ““New Jersey Mud 
Pits and the Glories of Fossil Insect Work,”’ 
presented by Dave Grimaldi of the Ameri- 
can Museum of Natural History, New York. 
Tickets are $24 per person, $12 for students 
and half-people. Dave Furth dominated the 
rest of the exhibits: notes from the ESW 
executive e-meeting, at which it was decid- 
ed that starting in October the regular meet- 


1092 


ings would be held at 7:00 pm; announced 
the BugFest on the Mall and again invited 
members to participate; announced the 
opening of a one-year exhibit on ‘‘Vanish- 
ing Pollinators” at the National Zoo in the 
Amazonia Science Gallery; displayed some 
new books: /nsects and Their Spermatozoa 
and Their Phylogeny; Catalog of the Het- 
eroptera of the Palearctic—Miridae;, The 
Preying Mantids; and The Butterflies of 
Hong Kong, which in addition to the usual 
photographs gives details on their mor- 
phology, larvae, and parasites. 

John Brown announced the evening’s 
speaker, Michael G. Pogue of the System- 
atic Entomology Laboratory, based in the 
National Museum of Natural History. He 
spoke to us about “The Armyworm Genus 
Spodoptera Guenée (Lepidoptera: Noctui- 
dae): a Multimedia Review.’ Spodoptera 
has 30 species and includes the single most 
serious pest of Lepidoptera; other species of 
the genus are also serious pests. The genus 
is mainly subtropical and tropical but some 
migrate into temperate regions. The genus 
is particularly abundant in Africa, Southeast 
Asia, Borneo, and South America. Dr. Po- 
gue’s monographic goals are to treat every 
species comprehensively, estimate their 
phylogeny, build a specimen database, and 
provide an interactive key. Features he uses 
to delimit species are taken from wing col- 
or, genitalia of both sexes, and larvae. The 
database actually consists of six linked da- 
tabases: specimens (more than 10,000 re- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


cords), a gazetteer (more than 4,000 Spo- 
doptera localities with latitude and longi- 
tude), larval hosts, parasitoids, predators, 
and a bibliography. The latter four are now 
available on the WWW (http://160.111.87. 
10:591/entomology/spodoptera/spodintro. 
html). The Expert System is in progress. 
Although primarily a key, the system is de- 
signed to give experts additional informa- 
tion once an identification is attained. The 
system uses the LucID windows program 
(University of Queensland and CSIRO), 
which is very flexible. The builder can code 
character states as polymorphic or “‘likely 
misinterpreted,’ options difficult to incor- 
porate into traditional, couplet-based keys. 
The user can choose characters in any order, 
display characters ranked by difficulty, and 
see existing images of any character states. 
Each choice produces a shorter list of pos- 
sible species. A traditional key for 30 spe- 
cies will have 29 couplets, so to identify the 
majority of the species a user passes 
through 15 couplets. In comparison, iden- 
tification paths using LucID are much 
shorter. For example, any species of Spo- 
doptera can usually be identified within five 
character state choices. Dr. Pogue plans to 
continue adding to the database as time per- 
mits. 

The meeting was adjourned at 8:55 pm. 
Refreshments were provided by John 
Brown. 


Respectfully submitted, 
Stuart H. McKamey 
Recording Secretary 


PROC. ENTOMOL. SOC. WASH. 
102(4), 2000, pp. 1093-1099 


PROCEEDINGS 
of the 
ENTOMOLOGICAL SOCIETY 
of 
WASHINGTON 


Volume 102 


OFFICERS FOR THE YEAR 2000 


President David G. Furth 
President-Elect John W. Brown 
Recording Secretary Stuart H. McKamey 
Corresponding Secretary Hollis B. Williams 
Treasurer Michael G. Pogue 
Program Chair Theodore R. Schultz 
Membership Chair Steven W. Lingafelter 
Custodian Jon A. Lewis 
Editor David R. Smith 
Past President Michael E. Schauff 


Published by The Society 
WASHINGTON, D.C. 
2000 


TABLE OF CONTENTS, VOLUME 102 


ARTICLES 
ABRAHAMSON, WARREN G.—See MELIKA, GEORGE ..........---...s ss eee cece sees cece tenes 198 
ADAMSKI, DAVID and ERIC H. METZLER—A new species of Glyphidocera Walsingham 
from southwestern Ohio (Lepidoptera: Gelechioidea: Glyphidoceridae) ..........-....-++..+++5 301 
ADLER. PETER H:i=—See PAYSEN, BRIG SF oiii es. s cece eee n cc cee es soe ce cm seecimneecimenenmerccs sc 843 
/NUICIQOIN|, ID) ——$"See TIE, VS Soke ssoncesosneoobosbenconasapeposaeccodanecogesossce0s0sec dosccoboc 688 
ALONSO-ZARAZAGA, MIGUEL A.—See HOEBEKE, E. RICHARD .............--...---++555 151 


ALUJA, MARTIN, JAIME PINERO, MAURILIO LOPEZ, CESAR RUIZ, ALBERTO ZUNIGA, 
ENRIQUE PIEDRA, FRANCISCO DIAZ-FLEISCHER, and JOHN SIVINSKI—New host plant 
and distribution records in Mexico for Anastrepha spp., Toxotrypana curvicauda Gerstacker, Rha- 
goletis zoqui Bush, Rhagoletis sp., and Hexacheta sp. (Diptera: Tephritidae) .........-.----++++++5 802 
ANTOLIK, CHRISTOPHER—See ROONEY, THOMAS P. .........--. 22-222 cece eee ee eee eees 308 
ARCHANGELSKY, MIGUEL—Immature stages of Neotropical Hydrophilidae (Coleoptera): 
Hydramara argentina (Knisch, 1925) and Hemiosus bruchi Knisch, 1924 ..................... 280 


1094 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ARCHANGELSKY, MIGUEL—See BRANHAM, MARC@ Av core ec ceene cece ccc saecueanes cece 869 
ASSIS-PUJOL, CRISTIANE VIEIRA DE and MICHEL LECOQ—Comparative study of sper- 
mathecae in eleven Rhammatocerus Saussure 1861 grasshopper species (Orthoptera: Acri- 


didae: Gomphocernae:S cyMlinini) hts 2 5cee en. dose ete ee eee An eee EEE en ae 120 
BAPTISTA, ALESSANDRA R. P. and WAYNE N. MATHIS—Notes on the genus Cyamops 

Melander (Diptera: Periscelididae), including description of ten new species .................. 481 
BARROWS: EDWARD INVE——Sceei CIC BRO MAR Ye Re meeea ee Cee er EEe etree eee te ere eEreaaee 50 
BLANCO:METZLER, HEEGA— See GAGNES RAYMOND S. spe.o502: sue se ee eee 831 
BORKENT, ART—The larva and pupa of Schizonyxhelea forattinii Wirth and Grogan (Diptera: 

Ceratopogonidae) with a discussion of the phylogenetic relationships of the genus ........... 862 
BOYDSTONWE==Sée PIRI, 6S (SP sccariccch tac ccn ademas Bese ee eee Taare A GER EEE eee ee 688 


BRANHAM, MARC A. and MIGUEL ARCHANGELSK Y—Description of the last larval in- 
star and pupa of Lucidota atra (G. A. Olivier 1790) (Coleoptera: Lampyridae), with a dis- 


cussion of abdominal segment homology across life stages. ...............0..cceeeeeceee cece sees 869 
BRASWELL, W. EVAN and JAMES R. OTT—The biology of Doa ampla (Grote) (Lepidop- 

tera: Doidae) on its host plant Stillingia texana (Euphorbiaceae) ..................... eee e eee e eee 507 
BRIGHT, DONALD E.—See VANDENBERG, NAWTATIA Jr o.oo oc ecenenesecccscseennene ceases seen 62 
BROWN, JOHN W.—Revision of Lobogenesis Razowski and Odonthalitus Razowski (Lepi- 

doptera: Tortricidae: Tortricinae), with comments on their monophyly .......................5. 72) 


BROWN, JOHN W. and JON LEWIS—Catalogue of the type specimens of Tortricidae (Lep- 
idoptera) in the collection of the National Museum of Natural History, Smithsonian Institu- 
trons WashinstomiyDNC), Rew rccieaececncomnh atane eaaika canis ensuu soa sass ac neeenias ance mertre meeen 1014 
BROWER, ANDREW V. Z.—On the validity of Heliconius tristero Brower Heliconius mel- 
pomene mocoa Brower, with notes on species concepts in Heliconius Kluk (Lepidoptera: 


IN Aion (0) 02 WG 21) oe ote are or ere tor en Seren Re Aer Senora Sac Penene ic nadonauondocaimaposcoe oC 678 
BUTLER, LINDA and JOHN STRAZANAC—Macrolepidopteran larvae sampled by tree bands 

in temperate mesic and xeric forests in eastern United States ................. 0.0 cece eee eee e ee eee 188 
BYERS ROBERT At Sees HORBERE, ES RICHARD a9. 2a. eee eee eee eee eee 151 
CICERO, MARY R. and EDWARD M. BARROWS—Milesine flower flies (Diptera: Syrphidae) 

in a central Appalachian broadleaf forest: Abundances, flight periods, and diflubenzuron. ........ 50 
CONTRERAS-RAMOS, ATILANO—A new species of Chloronia Banks (Megaloptera: Cor- 

ydalidae) from southeastern Brazil, with a key to the species of Brazil ........................ 919 


CRYAN, JASON R. and LEWIS L. DEITZ—Review of the New World treehopper tribe Ste- 
gaspidini (Hemiptera: Membracidae: Stegaspidinae): HI: Flexocentrus Goding, Stylocentrus 
Staleand"Wimibelligertus MD GitZs conc cece cass Se eee ae ee EERE ee 82 

DARSIE, RICHARD E, JR.—Description of the pupae of five species in the subgenus Armi- 
geres, genus Armigeres Theobald, with a key to species of the known pupae of the subgenus 
(Diptera GultcrGae) MRS x kee terre hse ese cid este ROS AT RSet ERE 108 

DARSIE, RICHARD EF, JR.—The pupae of three species in Minomyia Theobald from Nepal 
and a key to the known pupae of the genus in the Oriental and Australasian regions (Diptera: 

(GU LKec EVO) Seta meecmrcen ane aoc Gana accd mien ecu en EO mnan cr aae nom ane ma coaeommnnnboeranmacce noc 625 

DARSIE, RICHARD FEF, JR.—Description of the pupa of Armigeres (Leicesteeria) omissus 
(Edwards) and a key to the larvae and pupae of the Armigeres occurring in Nepal (Diptera: 


(SULT ARTO) eee ae ee aeRO et Berens at Seen EO Me See Shen | EPP ERE Aas taaaace cnosGoacdcoous 964 
DAVIDSON; JOHN ‘A.——See) POLAVAIR ABU Ss 7 ieet oecct ast cence otc car er senieee eee cereeeee 549 
DEIVZ, LEWISjL.—See'CRYAN, JASON Reiiks a.p.cbh asa aeaeton- usleceaaeae teahi ae eee eee 82 
DIAZ-FLEIS CHER. See ALUIA,, MARTIN. 4. cere oe eae cae x eee ee 802 
DIETRICH, C. H.—See NOVIKOV,.D! Vis .ns2Sohaaserens os teangctnteedan 6 15 setie deemee eee eee 170 
ECKERLIN, RALPH P. and HARRY E PAINTER—New records of fleas (Siphonaptera) from 

eastern West Virginia’ 3...) 00.60 cama nae sas gohan olaumiene rence maemisece eee eee eee ae eee eee 969 
EDMISTON, JAMES F——SeesMiATHITS SWAYINIESINets-se-e- oe ea deaeieece de ene he aea-eeeee eee 652 
EDMONSONER> E-=Scé) WALES ON AR ID Seer ea ee neo ee enolase och nee Lee eee ee 360 
ETIENNE SBAN —=SeeiGAGNEVRAY MOND: 205.254 s.o. cee ee oe ee 831 
EVANS, HOWARD E.—Three new species of Dipogon Fox (subgenus Dipogon) (Hymenop- 

tera: Pompilidae) from central and western North America ..................0.seeeee cece eres es 1010 
FOOLE, BENJAMIN A:——See NORRBOMEPAIBIER Noe aepe-peeee eee aae eee eee eee 142 


FOSTER, GEORGE A. and WAYNE N. MATHIS—Notes on Neotropical species of Tethina 
Haliday (Diptera: Tethimidae)) eee hn ee ae ae ees ees see aE 542 


VOLUME 102, NUMBER 4 


FROESCHNER, RICHARD C.—See HENRY, THOMAS J. ...... 2.02... c cece eect meee eset e eee 
GAGNE, RAYMOND J., HELGA BLANCO-METZLER, and JEAN ETIENNE—A new Neo- 
tropical species of Clinodiplosis (Diptera: Cecidomyiidae), an important new pest of culti- 
vated peppers (Capsicum spp.: Solanaceae) ............ 6. cece eee cece eee e teen eee eees 
GAIMARI, STEPHEN D. and MICHAEL E. IRWIN—Revision of the mexicana-group of the 
cycloteline genus Ozodiceromyia Bigot (Diptera: Therevidae) .................-s2. ee eeeee seen ees 
GATES, MICHAEL W.—A new species of Cirrospilus Westwood (Hymenoptera: Eulophidae) 
fromthe southwesterm United (States! and! MEXICO) foc ee cre crclete ore -iociatnie s)eisteialnla = ole eteteter= lores ete elelerotaa = 
GOEDEN, RICHARD D.—Life history and description of immature stages of Neaspilota sig- 
nifera (Coquillett) (Diptera: Tephritidae) on Hemizonia pungens (Hooker and Arnott) Torrey 
and A. Gray (Asteraceae) in southern Califormia .................2.:.eseseeseeeeeecerccssce estes 
GOEDEN, RICHARD D.—Life history and description of immature stages of Neaspilota aenig- 
ma Friedberg and Mathis (Diptera: Tephritidae) on Erigeron divergens Torrey and Gray 
(Asteraceae) mms outherns@alifomialpeeeeerreee eee eres ee tere eee neat eee pee eer rye er reer 
GOEDEN, RICHARD D.—Life history and description of immature stages of Neaspilota ap- 
pendiculata Freidberg and Mathis (Diptera: Tephritidae) on Machaeranthera canescens 
(Pursh) A. Gray (Asteraceae) in southern California ...................+.ssecceeneneecc seme ecerees 
GOEDEN, RICHARD D.—Life history and description of immature stages of Neaspilota pu- 
bescens Freidberg and Mathis (Diptera: Tephritidae) on Lessingia filaginifolia (Hooker and 
Arnott) M.A Wane (Asteraceae) ini southern Califormia cose ote ate ornins oa e)oele eialeieteletele oieiel= == 
(GIRN EMG = Ses THUMB KS Se) ohpnooranecasboasasnsaceA00 10060 9600000 cons conn apadaabauposspeeapodooonoce 
GUILBERT, ERIC—Revision of the genus Parada Horvath (Hemiptera: Tingidae) with cla- 
GUSTIN. AUTEN. 5 Ata aa Me cteiooic Secon OA BAAS aR oee tas SRS en ano esac So aCe oro cee Sessa ocr Sanaa: Oras 
EU ANIKS SIEAWIREINGE IME Sees MOORES ROBERT Greece cree eee eee reer ase eer: 
BUN RIBVACISE RUNILIPIEL 1, See IMO SITES. IDIOISIING ZN, 30254590000000 o0nosoerenodus sass ganoeaccasHonGs 
HASTRITER, MICHAEL W.—Jordanopsylla becki (Siphonaptera: Ctenophthalmidae), a new 
species of flea from the Nevada Test Site ............... 1... e ese e cece cece eect e eee ee tenes 
HASTRITER, MICHAEL W. and EUSTORGIO MENDEZ—A review of the flea genera Hec- 
topsylla Frauenfeld and Rhynchopsyllus Haller (Siphonaptera: Pulicidae) .............-....-.-- 
HENRY, THOMAS J. and RICHARD C. FROESCHNER—Corrections and additions to the “Cat- 
alog of the Stilt Bugs, or Berytidae, of the World (Insecta: Hemiptera: Heteroptera)” ............ 
HOEBEKE, E. RICHARD and A. G. WHEELER, JR.—Telmatophilus typhae (Fallén) (Cole- 
optera: Cryptophagidae), a Palearctic cattail specialist established in the Canadian maritime 
[DRO LE (CE SinGtilaeiee wigs cde OG REE DUS onc Hone En a Tne Peaa Sa RaA See TOS OC USSR OSS Ba REAR EenmEr ooo Praag P9aF 
HOEBEKE, E. RICHARD, ROBERT A. BYERS, MIGUEL A. ALONSO-ZARAZAGA, and 
JAMES E STIMMEL—/schnopterapion (Chlorapion) virens (Herbst) (Coleoptera: Curcu- 
lionoidea: Brentidae: Apioninae), a Palearctic clover pest new to North America: Recognition 
featuness GIStmbUtOMe and) DIOMOIMMIGS cece nee cesses eee elelelciclalars ciate le etet eye ole ter=l= ele) =Jelelalal-le(oleieroinietciei= 
HOWARD, THERESA M.—See MOSES, DUSTY A. ...............0s sees eect eee eee e cece cece 
HUNG, AKEY C. E and HACHIRO SHIMANUKI—Nucleotide sequence and restriction site anal- 
yses in three isolates of Kashmir bee virus from Apis mellifera L. (Hymenoptera: JN GES) sesooc 
HUSBAND, ROBERT W.—Redescription of Eutarsopolipus desani Cooreman and description 
of E. mirifica, n. sp. (Acari: Podapolipidae) from Chlaenius spp. (Coleoptera: Carabidae) 
MRO Sieur /NGOISEL Bon bb dduaoosoewosns boodacudeoon codeennanudoaunopodeonoTusno0es sesoHeD aude DpeeCdoUscoOs 
IRWIN, MICHAEL E.—See GAIMARI, STEPHEN D. ...........-- +--+ +--+ 22s eeeee teen tenet 
JAMESON, MARY LIZ—Synopsis of the Mexican and Guatemalan genera Rutelisca Bates 
and Metapachylus Bates (Coleoptera: Scarabaeidae: Rutelinae) with comments on classifi- 
cation of the subtribe Rutelina ................2..... cece eect eect ee etter cece tenet ees cc eee eeceecs 
JENSEN, ANDREW S.—Eight new species of Macrosiphum Passerini (Hemiptera: Aphididae) 
from western North America, with notes on four other poorly known species ..........-..--. 
JOHNSON, NORMAN F—See MUSETTI, LUCIANA ............... 2022222 s eee teen eee n eee 
KEIPER, J. B. and W. E. WALTON—Biology and immature stages of Ochrotrichia quadrispina 
Denning and Blickle (Trichoptera: Hydroptilidae), a spring-scraper .........-.++-+++eeeseeeeeee 
KIMSEY, LYNN S.—The western Australian genus Oncorhinothynnus Salter: New species and 
relationships (Hymenoptera: Tiphiidae: Thynninae) ........... 6... .60s sees eee e eee eee eet 
KOCH, FRANK and DAVID R. SMITH—Nematus oligospilus Forster (Hymenoptera: Ten- 
thredinidae), an introduced willow sawfly in the southern hemisphere ..............--...+++.++- 
KONSTANTINOV, ALEXANDER S.—See KOROTYAEV, BORIS A. ............------2222055: 


561 


69 


384 


519 


1003 


398 


178 


1096 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


KOROTYAEV, BORIS A., ALEXANDER S. KONSTANTINOV, and CHARLES W. 
O’ BRIEN—A new genus of the Orobitidinae and discussion of its relationships (Coleoptera: 


Curculionidae) siti 2 sasinbass date Ricken Stale = oie saeale Reet ete ASE to a ein eats eet eee O29 
IKROMBEIN, KARE RY.—_SceawAIs, RAYMOND ite eaeeeereceee ree atte cee Een re enaeee 271 
KRUSE, JAMES J.—Archips goyeriana, n. sp. (Lepidoptera: Tortricidae) an important pest of 

baldcypress (Taxodiaceae) in Louisiana and Mississippi ...............-.....ceeeeeeccceeeseeceaes 741 
LAPIERRE, LOUIS M.—Prey selection and diurnal activity of Holcocephala oculata (F.) (Dip- 

tera: “Asilidae) in | COStamRiGa rab osticb Soo ae eA eae Ee ne Ae Pe RE 643 
LECOOs MICHELE See ASSIS-PUIOENERISTIANESVIER AIDE a eee ne ane een eee re 120 
LEMMON CAROTR==sees MATER. GCHRISME Wieen eee eeeeee ee eee eee aeaee ae aes 747 
EESTER;, DiGi =—SeesWwileSON: Az Di iin d5ccdten cc Ee, SE A nee 360 
BEWIS; VON ==SceeiBiRO  WNI ORIN OWS  eeiecnccsncescocne sae rises ceereniosaceionn i biarirosactee ie aan 1014 
LOPEZ MAURIIO™=s ce FATWA SIAR TING: Oe eC ee ee ee 802 
LOPEZ-MARTINEZ, VICTOR—See WHARTON, ROBERT A. ............0:cc0secceeeeeeeereees 794 


MAIER, CHRIS T. and CAROL R. LEMMON—Discovery of the small Japanese cedar lon- 
ghorned beetle, Callidellum rufipenne (Motschulsky) (Coleoptera: Cerambycidae), in live 


arborvitaemn"Gonnecticut™ 2: A dtace lane ones nen one fone beac ewer sete mgt banades ae eenres son aeenre anaes 747 
MARSHALL, S. A.—Chespiritos, a new genus of Limosininae (Diptera: Sphaeroceridae) from 

GOStAMRIC AM en. sass eso: ote as Stee Seele eee eae EEE ce coher E ae Sein eetes Hon ae SoM an emer me ea 609 
MATHIS, WAYNE N.—See BAPTISTA, ALESSANDRA R.P. 122.2202... 22sec. ccccceesecceee esse 481 
MIAUENIS SWAIN INS ees OSMERS GEORGE TA ec saeco eee eae ener een ae ee eeeee cn eeee 542 
MATHIS, WAYNE N. and JAMES EF EDMISTON—A revision of the shore-fly genus Lem- 

naphilat@ressony(Dipteray Ee phy dndac)t-.-eacce ace see sec eecis see eee e ee eee eee eer 652 


McKAMEY, STUART H.—A review of the New World leafhopper subgenus Texananus (lo- 
wanus Ball) (Hemiptera: Cicadellidae: Deltocephalinae) with a checklist and distribution sum- 


IMAaby 1Oh SPEClESMM te GENUS acne eneren cen on tren omar connie etnies ame tm seria eee aeienen eer DN 
MELIKA, GEORGE and WARREN G. ABRAHAMSON—Review of the cynipid gall wasps 

of the genus Loxaulus Mayr (Hymenoptera: Cynipidae) with descriptions of new species... 198 
MENDEZ, EUSTORGIO—See HASTRITER, MICHAEL W. .-...0..-.5-c0..)21s cess eee 613 
METZEBR. ERIC Hi See ADAMSRIn DA VID en nc se seecesnprecls see Sec ae Aee Oe ace eee Seen 301 
MICE RS DOUGEASS R——see ROLAVARARUS Saeccaeise ee rec ee eee ener eee eee eeeree 549 
IVIIGTE EAR RE aS CO PUIG RS ca crane ee rsiacerctea Menara eactaltevarcias ie se svn seg hue ePeveas acefol ae orate eI ee ae sea eee ORE 688 
TAA OU Ce) EH Saeed eS ereiel 2 UI Sa ahs aes Noein a nearer arene seacmangaedaacdcade suodanoseroeanacoos 688 
MOORE, ROBERT G. and LAWRENCE M. HANKS—Avian predation of the evergreen bag- 

worm (kepidopteta: ESVChidae) Gana waeracerreeeeteee ace n ore cere eeeCe eer ee Peete eens 350 
IMMORVAING, IMUANIMIU SUEY IDK =e IROLOINDE NG INSKOIMUNS) 12. Gonasccoocadooenscnosononovcsdaadedcanceecs 308 
MORON, MIGUEL ANGEL and ANGEL SOLIS—New Costa Rican species of Phyllophaga 

Flarnish (Coleoptera Melolonthidac-)Melolonthinae) saa-eere eee eeeee eee ere een eee cree eee 901 
MOSES, DUSTY A., THERESA M. HOWARD, and RALPH E. HARBACH—A new species 

of the subgenus Sabethoides of Sabethes (Diptera: Culicidae) from Venezuela and Brazil .. 991 
MUSETTI, LUCIANA and NORMAN FE JOHNSON—First documented record of Monomachidae 

(Hymenoptera: Proctotrupoidea) in New Guinea, and description of two new species ............ QSi7/ 
NEUNZIG, H. H.—Uncitruncata leuschneri, a new genus and species of Phycitinae (Lepidop- 

teraweyralidac) proms Calitoniarandi@Ores oniee-eeeeer eee eee eee eerGeereeeet eee Eee eee ens 408 
NEUNZIG, H. H.—New species of Puerto Rican Phycitinae (Lepidoptera: Pyralidae) ......... 838 
NORRBOM, ALLEN L. and BENJAMIN A. FOOTE—A new subgenus, Footerellia, and new 

distribution records of Neaspilota Osten Sacken (Diptera: Tephritidae: Terelliini) ............ 142 
NOVELO-GUTIERREZ, RODOLFO—Description of the larva of Hetaerina infecta Calvert 

(Odonata: Calopteryeidae)) (o.xtee de saswjac enue. qeae-erre ne discs <ckn-oa scene seem tere eer ee 99 
NOVIKOV, D. V. and C. H. DIETRICH—New megophthalmine leafhoppers (Homoptera: Ci- 

cadellidae) strom) Mexico; withiaskeystosNews Worldsspeciesiys..--ems-teeteaseeeee eee eee eee 170 
© BRIENT CHARLES W:—seesKORODMYAEV, BORIS “Ag fe) sepa eae ee eee eee ee ere eee 929 
OTT, JAMES R:— Sees BRASWELL AW HENAN) \.c:restecerocen tease ae ere eacee eeecie: eoeeecteene 507 
PAINTER, HARRY F——See BEMERIEINGRVAISP He Psa eee eee acne eee a eee eee 969 


PAPP, JENO and SCOTT R. SHAW—A study of the genus Falcosyntretus Tobias from the 
New World with five new species and a key to known species (Hymenoptera: Braconidae: 
Euphorinae)) s,s.scsea stipe cbse ee Se ee ae EE Sa ee Eee 634 


VOLUME 102, NUMBER 4 1097 


PAYSEN, ERIC S. and PETER H. ADLER—Taxonomy and polytene chromosomes of Simu- 


lium parnassum Malloch (Diptera: Simuliidae) .................6.20 ss eee cee cece ee cee eee eee e cee ees 843 
PEDROSA-MACEDO, J. H.—Biology and behavior of the strawberry guava sawfly, Haplos- 

tegus epimelas Konow 1901 (Hymenoptera: Pergidae), in southern Brazil ...............-....- 129 
PIEDRASENRIOUE—See.A EUSANIARIBING)..2:: <aib eae deniers neat eee ee 802 
PIKE, K. S., P STARY, T. MILLER, G. GRAF D. ALLISON, L. BOYDSTON, and R. MILL- 

ER—Aphid parasitoids (Hymenoptera: Braconidae: Aphidiinae) of northwest USA .......... 688 
EINERO.VAIME See AWUTA: MARTIN: o6.0:.c8h stasis tie sel Ghee TE vrs an de ee 802 
POGUESMICHAE LI G:——Sceer SALUKES SANDRAOWV. co ie cen crecena-eeeteceoce caste ernneer aa ee 233 


POLAVARAPU, S., JOHN A. DAVIDSON, and DOUGLASS R. MILLER—Life history of 
the Putnam scale, Diaspidiotus ancylus (Putnam) (Hemiptera: Coccoidea: Diaspididae) on 
blueberries (Vaccinium corymbosum, Ericaceae) in New Jersey, with a world list of scale 


INSECtSKONMDIMEDE RGLES reenact oer ers Oras 2 nue mee ealoiQak ale aiere lays clenals Measeielbvare stato Ne erstelattstee 549 
POLHEMUS, DAN A.—A revision of the endemic Hawaiian reduviid genus Saicella Usinger, 

with descriptions of four new species (Heteroptera: Reduviidae: Emesinae) ................--. 1 
QIAO, GE XIA and GUANG XUE ZHANG—A taxonomic review of the genus Delphiniobium 

Mordvilke,(@iemeptera: Aphididae)an (Chimay S722 censen eee 222 eee eierolenemeleeinccine reser 892 
RABAGLIA, ROBERT J.—See VANDENBERG, NATALIA J. 22.2.0... 5.000222 ce ccc cece esse cece 62 
ROBBINS, ROBERT K.—The New World hairstreak genus Arawacus Kaye (Lepidoptera: Ly- 

CASMCESS INocelteies IK SN) pone sogd>quacodsenodeseacone s4d0e ce cetacodenouaeabsenopsopscopadoasun 162 
ROONEY, THOMAS P., CHRISTOPHER ANTOLIK, and MATTHEW D. MORAN—The im- 

pact of salamander predation on Collembola abundance ................... 222s. see eeeeee eee ee ees 308 
RIZE SAR See VAT UNA INMANROTAING 0 bat tee ans encr eres eetean eed atcae 3 are ae ieee one 802 


RUSSELL, LOUISE M.—Notes on the family Aleyrodidae and it subfamilies: Redescription 
of the genus Aleurocybotus Quaintance and Baker and description of Vasdavidius, a new 


genus (Homoptera: Allyerodidae) 2.2. vce recimersiecins seem cles ore femelle ole elses sieve vis tine nice 374 
S/ATINIL IMUAILIRIUNIT S—=Gyae, WANSIUR YG cacacoucaocsndosoncogaqoodasdndssepene coon oosnRooNcoD doo BasdcoORC 601 
SALUKE, SANDRA V. and MICHAEL G. POGUE—Resolution of the Elaphria festivoides 

(Guenée)) species complex (Lepidoptera: Noctuidae) ........... 02. iiii eee eee ee eee 233 


SCARBROUGH, A. G.—Two additional species of robber flies of the genus Ommatius Wie- 
demann (Diptera: Asilidae) from the Bahamas and with replacement names for two other 


5) DSLETER) 5 ce done OS RIBIe ae 9 SORTA Cre SDD A FB RES AR TRCADO dre Gh Ec Loc ge dap. 7 cGE RO OnOCanA gp aaa opedoee oF 912 
SCHAUFE MICHAEL E.—A new genus and species of Eulophidae (Hymenoptera) from Costa 

RUA Symlilal TMOWES Cin HS SSNS, 0450000 cosdocssacnedeasesseboeunD on CSusscauuODSUdODeodoHseacAboAneauacac 403 
SHAWASCO IE IR: —See- PARP? JENO 6 .x).5 ccc cea ren scl ents te oo nosed dation athe Aevaetes HeseaO ae 634 
SHIMIANUIKG. HA CHIR @— See EIUINGay AIK (© isteresereteererreretete toler s)o ro = eroded =letetel- fatale tate tte) 178 
SITES, ROBERT W. and MICHAEL R. WILLIG—Morphometric variation among populations 

of Ambrysus mormon Montandon (Heteroptera: Naucoridae) .............-.-+++see esses sees eee 533 
SIVINSKLVIOHN= See AlUNAs MARTIN, 35.case¥ eer nets eS ele SR 802 
SIMIMOSE IDVAUD) IRi——Syee IM(OLEI8 F JERVAINILG Go cooduasessncocenencnesanees coodbbenHSbaccseodnsacace ono 292 
SIMICMaL IDZAWALD) IR Seo WANSIOR WG ccvosccaconscsb0osonccns caus eon bocGndnocunoposeccocdonooToTecRosont 601 
SIMIGTEEs AWAD RES ee) ZIIN @) VIE VSPA EGG Gos Freee eters erste evoke ele rele oleate let dedeleke fale let elated =r stalet=tele 852 
SIMMS E IDYANA0D) Re Srees ZAIN(O UIE, ZENE EN'C (Gi oa6 GondacaasenaccadoopuaoddeSdencspsecescnuancc 974 
SOLIS S ANGEL — See: MORON) MIGUEL 2ANGED iwc. .cacyah sen sens she ete thence ones aeincm tener 901 
SPIRO GP" See DIK Ke. Sn 8 A or eerye rete isi: eee. (Sauda Sacto coe eee 688 
STARVAZZ/NINUNGS JOVSIN Ses IBIUMNLIENRS ILINIDYA 5 ocooncse0ccne0sconbeoongrogscodogpeuadnavacnssesabnan 188 
SEIMMETDT, JAMES! E—See HOEBEKE, Es RICHARD) Jo 5225. cae -- acters lee =e ese ame 151 
THOMPSON, FE CHRISTIAN and MANUEL A. ZUMBADO—Flower flies of the subgenus 

Octyptamus (Mimocalla Hull) (Diptera: Syrphidae) ............. 22... eee eeee eee eee ee eee eee e eens V3 
TOGASHI, ICHIJI—Description of a new species of the genus Stiricorsia Konow (Hymenop- 

ieee OSS Ch) ison EVEN Soocscuoscnnnaenseadobo0bpoo Sb occduonEscenasuddaSsseoaesAboconAen ooasqeocc 105 
TOGASHI, ICHIHI—Japanese sawflies of the japonicus group of the genus Taxonus Hartig 

(Hymenoptera: Tenthredinidae) .................. ee eeeeeeee eee eee eee e eee eee een e eee eeenencees 33 
VANDENBERG, NATALIA J., ROBERT J. RABAGLIA, and DONALD E. BRIGHT—New 

records of two Xyleborus (Coleoptera: Scolytidae) in North America ..........--..+-.+00e005- 62 
VASU, V., DAVID R. SMITH, and MALKIAT S. SAINI—Review of the Asian sawfly genus 

Anisoarthra Cameron (Hymenoptera: Tenthredinidae) .................+. 22s eee eee e reese eee eee 601 


WAHIS, RAYMOND and KARL V. KROMBEIN—A new Machaerothrix Haupt from Sri 
Lanka with notes on the genus (Hymenoptera: Pompilidae: Pepsinae: Ageniellini) ........... NWA 


1098 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


WALTON WB: Seer RBIPBRe 7 Bit Sion. gaiacas tn dene sete som neers oooh TL an ener 183 
WELLS, SAMUEL A.—Two new species of Horistonotus Candeze (Coleoptera: Elateridae), 
new synonymies, and a key to the species of the United States and Canada .................. 412 


WHARTON, ROBERT A. and VICTOR LOPEZ-MARTINEZ—A new species of Triaspis 
Haliday (Hymenoptera: Braconidae) parasitic on the pepper weevil, Anthonomus eugenii 


Canox(ColeopteraxGurculionidae)) Seco. secon eect lee ee Sar ren RES eRe SEERA eee eee 794 
WHEBRER (ASG; JRe— Sees HOEBE KEE MRICHARD Ieeenaseen eee nae ee eee eee eee 398 
WIETETG VMIGHABIOR Sees iES. ROBE RU Wiareesseaeeaseeeeeee een ehhe eee eee eeeeee 5535 
WILSON, A. D., D. G. LESTER, and R. E. EDMONSON—Live oaks, new hosts for Odon- 

tocynips nebulosa Kieffer (Hymenoptera: Cynipidae) in North America ....................... 360 
WOODLEY, NORMAN E.—A new Afrotropical species of Allognosta Osten Sacken (Diptera: 

S rat Onnyii Gd aAS) Per ata cer es ee a Se ne eee 924 
YASUNAGA, TOMOHIDE—Anthocorid bugs of the tribe Oriini (Heteroptera: Anthocoridae) 

of the /@Ocasiwarai(Bonin) islands Japaniytes.t a2 Sige a Bo, ae Paes 353 
ZEHANGIGUANGEXUE—=Scel@ VA ON GEUXTAG Wyse eee ae ee one en eae 892 
ZINOVJEV, ALEXEY G. and DAVID R. SMITH—Types of sawflies described in the genus 

Pontania A. Costa (Hymenoptera: Tenthredinidae) in the Illinois Natural History Survey ... 852 
ZINOVJEV, ALEXEY G. and DAVID R. SMITH—Sawflies (Hymenoptera: Tenthredinidae) 

described by Benjamin D. Walsh, with notes on their hosts and biology .....................+5 974 
ZUMBADO; MANUEBIE A See DHOMPSON, EF GHRISTIAN 252220) cee ee eee eee a3 
ZUINIGA:-Al-BERTO=See AWUIA. MARTIN 2:4,20 125.0 ERO ott ea eee 802 

NOTES 
ALUJA, MARTIN, EMANUEL HERRERA, MAURILIO LOPEZ, and JOHN SIVINSKI—First 

host plant and parasitoid record for Anastrepha spatulata Stone (Diptera: Tephritidae) ...... 1072 
COCA-ABIA,, MILAGROS-—“See GARCIA=PARIS, MARION... 42 Len alse oe 473 
GARCIA-PARIS, MARIO, GABRIELA PARRA-OLEA, and MILAGROS COCA-ABIA— 

First records of the genus Noserus LeConte (Coleoptera: Zopheridae) in Mexico ............. 473 
IRI BININISRVAS JaIMUANNIOI BU Syeret JNILIUIUAS, IMIR, soca nagocs 900 asoovanopbvconcancooonauocenncscocass 1072 
JACOBS, EWGENE Ay——See FARTS TEPHENL IS csctvantatass seheconsaioaehiereee neon eee eae 75 
KERSTNER ;CORENNAD==Seev PAPEL) STEPHEN: laus.cciee: Maeda aecee eet ee eee eee 2D 
LAZZARI, SONIA MARIA NOEMBERG—See ZONATA DE CARVALHO, REGINA ...... 757 
LOPEZ, MARUILIO “See /AbUIAM MAR TING 2 hace 25.5528 s eee en ee 1072 
PARRA-OLEA, GABRIELA—See GARCIA-PARIS, MARIO) .....5.0..¢0.5.0005b2eccsc teehee estes 473 
PEASE VENGG—SceROBBINS RICHARDG A fest asta e ee eer cee renee 225 


ROBBINS, RICHARD G. and STEVEN G. PLATT—First report of Amblyomma clypeolatum 
Neumann (Acari: [xodida: Ixodidae) from the Union of Myanmar, with two new records from 


LOMEOISES 5258 stains eitee tha erie Seis ok by a arene enter ele eS a So Ee ee RR oes on St A eae 225 
SIVINSKE: JOHINSee- AL UIA. MARTIN 555m tise ovo nation lise PASO Ee AOR ee rae 1072 
TAFT, STEPHEN J., CORENNA D. KERSTNER, and EUGENE A. JACOBS—Ectoparasitic 

insects from migrating saw-whet owls (Aegolius acadicus) in central Wisconsin ............. IS 
WHEELER, A. G., JR.—New distributional and first specific host-plant records for Thionia 

acuta Doering and T. producta Van Duzee (Auchenorrhyncha: Fulgoroidea: Issidae) ........ 759 
WHEELER, A.G., JR.—Fitchia aptera Stal (Hemiptera: Reduviidae): Seasonal history and 

habitsan: mid-Appalachian:shale: barrens)... kasssanene eh ose eee oe ee eee eee 1070 


ZONTA DE CARVALHO, REGINA and SONIA MARIA NOEMBERG LASSARI—First re- 
cord of the California pine needle aphid, Essigella (Essigella) californica (Essig) (Homoptera: 
Aphicidacwleachninae) sine SOUtherny nazar e eee eres ae nee eee US 


BOOK REVIEWS 


BROWN, JOHN W.—Classification of Lepidoptera. Part 1. Introduction, by John B. Heppner ... 1075 
GAGNE, RAYMOND, J.—ldentification of North American Porricondyline Larvae (Diptera: 
Cecidomytidaamiby: Jolin’ D;, Plakidas: 17. cee ee asa ce Leon Ae eee eee 227 
ROBBINS, RICHARD G.—The Genus Rhipicephalus (Acari, Ixodidae): A Guide to the Brown 
Ticks of the World, by Jane B. Walker, James E. Keirans, and Ivan G. Horak ................ 768 


VOLUME 102, NUMBER 4 1099 


SMITH, DAVID R.—Pflanzenwespen Deutschlands (Hymenoptera, Symphyta) Kommentierte 


Bestandsaufnahme, edited by Andreas Taeger and Stephan M. Blank .......................... 229 
STOETZEL, MANYA B.—Aphids on the World’s Crops, An Identification and Information 
Guide, Second Edition, by R. L. Blackman and V. FE Eastop ..................0cce cece cece eeees 1074 
OBITUARIES 
ARNE TM ROSS) He JR —SeerS REINER] WARRENGES DIRe sacceesceereceee seen cere esas 761 
DETEZS BE WIS le —See*PARRIERS MAURICE, He 325 coccncekan satecen oo os tiies tate sis meiseesaulsieer ec 1077 
FARRIER, MAURICE H. and LEWIS L. DEITZ—Clyde Fuhriman Smith (1913-2000) ...... 1077 
IKINGSOLVIERSJOHNeME——See Ss MEINERS WARREN) Ex IRQ aaaecseetecee cea ote sei see errr 761 


STEINER, WARREN E., JR., JOHN M. KINGSOLVER, and ROSS H. ARNETT, JR.—A 


HRLOLTNES (1) eral SyovllonBn SooseeeaconeccasncoosososocsoubogsgnGpnaadobo0ccpene spobogococcosDEsoDDLSNEDdOC 761 
MISCELLANEOUS 

INSTRU A CONS Itoe GINO O MT ese peobooas cone cis Gu ocodndedonon naa ES pa noonRerH EBA ceMunnanpecnnrocmcocaactisa 771 

i eeO RG Cit ONES RS Geeennanepecns csanedo sue cooc ecb sorcontncr CoEBe Hest tae or enn aan scab 1c Oro dcEr ocr 475 

SOS IRIN SSIES sacra aosipon7 ase iecee cbce nooo eegoncan Stes sce uo aa oU sEac acerca spncedcnn asa 477, 1088 


EAblero tl GOntents sn, O LUTE wil De praceee ariel. esa etaeva rate rats erate No cde arralcvo id ioe ce aIsre eee eter Sevare eeieerareiets 1093 


1100 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
Entomological Society of Washington 


Publications.—Available Miscellaneous Publications and Memoirs of the Society are 
listed on the inside back cover of each issue. 


Instructions for Authors.—Instructions were last published in 2000, 102(3): 771-772. 


Bylaws.—The bylaws were last published in 1966, 98(3): 610-614. Several amend- 
ments were proposed at the December 3, 1998 regular meeting and passed at the January 
7, 1999 regular meeting. See minutes of the Society meetings, 1999, 101(2): 467 (1034th 
regular meeting), and 1999, 101(4): 920 (1035th regular meeting). 


Membership.—A membership list is published every three years and was last published 
in 1999, 101(4): 914-919. 


Seal.—The history of the official seal of the Society was published by Jon L. Herring, 
1964, 66(1): 1, and was reprinted in 1997, 99(1): 208. 


History.—The following articles relate to the history of the Society: 


Wade, J. S. 1936. The officers of our Society for fifty years (1884—1934). Proceedings 
of the Entomological Society of Washington 39: 121-132. 

Gurney, A. B. 1976. A short history of the Entomological Society of Washington. 
Proceedings of the Entomological Society of Washington 78(3): 225—239. 

Spilman, T. J. 1984. Vignettes of 100 years of the Entomological Society of Washington. 
Proceedings of the Entomological Society of Washington 86(1): 1—10. 

Stoetzel, M. B. 1984. ESW past-presidents for the years 1884 through 1983, photo- 
graphs and support officers. Proceedings of the Entomological Society of Wash- 
ington 86(1): 11-35. 


The papers by Spilman and Stoetzel were part of the centennial issue of the Society, 
1984, 86(1). The centennial banquet was held at the University of Maryland, College 
Park, March 12, 1984; see Society Meetings, 1984, 86(4): 975-976. 


The 100th regular meeting of the Society was celebrated February 2, 1995 and was 
held at the Beltsville Agricultural Research Center, Beltsville, MD. It was attended by 
115 members and guests and written up in the minutes of Society meetings 1995, 97(4): 
897-898. 


PUBLICATIONS FOR SALE BY THE 
ENTOMOLOGICAL SOCIETY OF WASHINGTON 


MISCELLANEOUS PUBLICATIONS 


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 


Grissell and Michael E. Schauff. 87 pp. 1997 


MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Memoirs 2, 3, 7, 9, 10, 11, and 13 are no longer available. 


No. 1. 


No. 21. 


No. 22. 


The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 ___.........- 
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 
EEE TE TSP Es) UEP CT oR 120) I Maa ACE eh aN ale oe eM aA Shes MGM A eek STONE Os SING BL 


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 _ 


. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. 


OOS GMS NES elk AST Se Sg alae, 22 AMS BCU A aM AMO CA Dea Ry. PN «COR Co BUPA Eira as 


. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174 


TTS te Rk ERS ag ES ENGR cele) a A Se a LON | AOL aI eal es eT, SE SSAA SATE 


. A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent 


CUR Ags Wig Soy ee Pa Pa So CS fo ca Re RP RRS, PST CARN EH iE See Ne Zane 


. 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 


ys. 2_ Norden. and) ALS Menke (21 G'ps: 1096 eal Be ae A he ra 


. Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and 


Pyatiaganle: (srogan, Jn 2Onppr hoo hi ets ek ee kU aE Trek okt) ae nah 


. Monograph of the Stilt Bugs, or Berytidae (Heteroptera), of the Western Hemisphere, by 


Noistnas:) benty C40 psi QO | eat ake BRO Ia pn de re ce! Ree 


. The Genera of Elaphidiini Thomson 1864 (Coleoptera: Cerambycidae), by Steven W. Lin- 


“SLT S Egy RYT ea IED? RR ee eee oO COOMERA ar a ALPEN a AIDS a Stemi MAR RM as ban ir 


New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae), by David G. 
Jay Tae UN Mao Bid LS 2: Sian A RRL Seep cee ae a ane) ae al We AER. FORGO, SQ eS UEC SAR teal EE aN som, Sdn Le 


Systematics of the North American Species of Trichogramma Westwood (Hymenoptera: 
iiachosrammatidae)sby John) Pinto 287. pp. L999 sane ae ee eee 


$10.00 


15.00 


$15.00 
15.00 
15.00 


15.00 


12.00 


5.00 


25.00 


25.00 
25.00 


25.00 


25.00 


18.00 


12.00 


12.00 


28.00 


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 
percent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the 
Custodian, Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 
Washington, D.C. 20560-0168. 


CONTENTS 


(Continued from front cover) 
+ 
HENRY, THOMAS J. and RICHARD C. FROESCHNER—Corrections and additions to the 
“Catalog of the Stilt Bugs, or Berytidae, of the World (Insecta: Hemiptera: Heteroptera)” .. 


KOROTYAEV, BORIS A., ALEXANDER S. KONSTANTINOV, and CHARLES W. O’BRIEN— 
A new genus of the Orobitidinae and discussion of its relationships (Coleoptera: Curculi- 
onidae) 


MORON, MIGUEL ANGEL and ANGEL SOL{S—New Costa Rican species of Phyllophaga 
Harris (Coleoptera: Melolonthidae: Melolonthinae) 


MOSES, DUSTY A., THERESA M. HOWARD, and RALPH E. HARBACH—A new species of 
the subgenus Sabethoides of Sabethes (Diptera: Culicidae) from Venezuela and Brazil 


MUSETTI, LUCIANA and NORMAN F. JOHNSON—First documented record of Monomachidae 
(Hymenoptera: Proctotrupoidea) in New Guinea, and description of two new species 
NEUNZIG, H. H.—New species of Puerto Rican Phycitinae (Lepidoptera: Pyralidae) 


PAYSEN, ERIC S. and PETER H. ADLER—Taxonomy and polytene chromosomes of Simulium 
parnassum Malloch (Diptera: Simuliidae) 


QIAO, GE XIA and GUANG XUE ZHANG—A taxonomic review of the genus Delphiniobium 
Mordvilko (Homoptera: Aphididae) in China 


SCARBROUGH, A. G.—Two additional species of robber flies of the genus Ommatius Wiedemann 
(Diptera: Asilidae) from the Bahamas and with replacement names for two other species .... 


THOMPSON, F. CHRISTIAN and MANUEL A. ZUMBADO—Flower flies of the subgenus 
Ocyptamus (Mimocalla Hull) (Diptera: Syrphidae) 


WHARTON, ROBERT A. and VICTOR LOPEZ-MARTINEZ—A new species of Triaspis Haliday 
(Hymenoptera: Braconidae) parasitic on the pepper weevil, Anthonomus eugenii Cano (Cole- 
optera: Curculionidae) 


WOODLEY, NORMAN E.—A new Afrotropical species of Allognosta Osten Sacken (Diptera: 
Stratiomyidae) 


ZINOVJEV, ALEXEY G. and DAVID R. SMITH—Types of sawflies described in the genus 
Pontania A. Costa (Hymenoptera: Tenthredinidae) in the Illinois Natural History Survey .... 


ZINOVJEV, ALEXEY G. and DAVID R. SMITH—Sawflies (Hymenoptera: Tenthredinidae) de- 
scribed by Benjamin D. Walsh, with notes on their hosts and biology 


NOTES 


ALUJA, MARTIN, EMANUEL HERRERA, MAURILIO LOPEZ, and JOHN SIVINSKI— 
First host plant and parasitoid record for Anastrepha spatulata Stone (Diptera: Tephritidae) .... 


WHEELER, A. G., JR.—Fitchia aptera Stal (Hemiptera: Reduviidae): Seasonal history and habits 
in mid-Appalachian shale barrens 


BOOK REVIEWS 
BROWN, JOHN W.—-Classification of Lepidoptera. Part 1. Introduction, by John B. Heppner .... 


STOETZEL, MANYA B.—4phids on the World's Crops, An Identification and Information Guide, 
Second Edition, by R. L. Blackman and V. F. Eastop 


OBITUARY 

FARRIER, MAURICE H. and LEWIS L. DEITZ—Clyde Fuhriman Smith (1913-2000) 
MISCELLANEOUS 

Society Meetings 

Table of Contents, Volume 102 


» Sve ke 


APRIL 01 


I 


l| 


| 


| 


| 


| 


| 


| 


|| 


88 00933 5670 


| 


| 


90 


Epes 


TA aees, 


Op ae ny ea eh 
treme be 


rotere 


wie ape 
esges 


AKA geome 
Spa areras 


Lae ap PE Nees ns 


preeern 


Puasa ive 
hen 


“Asa ee 


Nae